B74 
191 


55 


IN  MEMORY 

of 

Dr.   Clement  A,  Whiting 

PRESENTED  BY 

Dr.  Lillian  Burlier 


FIG.  A. — Spirochaeta  pallida  and  Spirochaeta  refringens.  Smear 
from  a  primary  syphilitic  sore,  fixed  in  osmic  vapour  and  stained  by 
Giemsa's  method  (leucocytes  necessarily  overstained) . 


I 


FIG.  B. — Spirochaeta  pallida  in  liver  of  syphilitic  foetus.     Stained  by 
Levaditi's  silver  method. 


UBF/iFY  OF 

COLLEGE  op  GG 
F  H  Y  s  f  r,f  A  f/s  J&  r 


I  ROCHET  ES3 


,4  REVIEW  OF  RECENT  WORK- 

ORIGINAL  OBSERVATIONS 


I.  BY 

W7  CECIL  BOSANQUET,  M.  A.,  M.  D. 

FELLOW  OP  THE  ROYAL  COLLEGE  OF  PHYSICIANS.  LONDON; 
FORMERLY  FELLOW  OF  NEW  COLLEGE,  OXFORD 


ILLUSTRATED 


PHILADELPHIA   AND  LONDON 

W.  B.  SAUNDERS  COMPANY 

1911 


*UU 

O:n\<0    nO  303JJOO 
i/;KMcM-iS 


Copyright,  1911,  by  W.  B.  Saunders  Company 


PRINTED     IN     AMERICA 


PREFACE 


HAVING  had  occasion  to  study  the  literature  bearing 
on  the  subject  of  Spirochaetes  in  connection  with  obser- 
vations which  I  made  on  two  species  of  this  genus,  I 
was  struck  with  the  voluminous  and  at  the  same  time 
scattered  mass  of  papers  to  which  reference  had  to 
be  made.  I  therefore  thought  it  might  be  useful  to 
others  if,  instead  of  merely  laying  aside  the  abstracts 
which  I  had  made,  I  put  them  together  in  the  form  of  a 
small  book. 

The  Spirochaetes  present  a  problem  of  considerable 
interest.  Their  position  in  the  scale  of  living  things  is 
still  undecided,  and  there  is  a  tendency  to  place  them 
in  a  position  between  the  bacteria  and  the  protozoa. 
My  own  observations  would  lead  me  to  regard  them  as 
much  more  closely  allied  to  the  former  than  to  the  latter. 

I  have  endeavored,  in  the  first  part  of  this  work,  to 
compare  the  statements  of  different  writers  on  the 
morphology  and  behavior  of  Spirochaetes,  with  a  view 
to  establishing,  if  possible,  the  features  characteristic 
of  the  genus.  In  the  second  part,  I  have  dealt  with  the 
separate  species  described.  I  have  confined  myself 
to  very  brief  descriptions  of  the  various  organisms  and 
of  the  methods  employed  for  examining  them,  as  those 
who  are  interested  in  particular  species  or  methods  will 
in  any  case  refer  to  the  original  papers.  Their  task 
will,  I  hope,  be  facilitated  by  the  bibliography  here 
appended.  This  is  by  no  means  exhaustive,  as  many 
papers  are  published  in  unattainable  periodicals,  and  I 
5 


6  PREFACE. 

have  included  comparatively  few  references  to  articles 
which  I  have  not  myself  consulted  either  in  the  original 
or  in  abstract.  I  hope,  however,  that  the  more  impor- 
tant papers  are  included. 

My  thanks  are  due  to  Mr.  A.  N.  Leathern  for  the 
loan  of  specimens,  and  to  Professor  E.  A.  Minchin  for 
the  very  kind  loan  of  books  from  his  library  and  of 
specimens  in  his  possession,  as  well  as  for  permission 
to  use  his  laboratory  for  my  own  small  researches. 

LONDON,  ENGLAND,  W.  C.  B. 

May,  1911. 


CONTENTS. 


SECTION  I. 
GENERAL  CHARACTERS  OF  SPIROCH^ETES. 

PAGB 

BIOLOGICAL  POSITION  OF  SPIROCH^ETES -    .    .  n 

MORPHOLOGY  OF  SPIROCH^TES 16 

Large  Spirochaetes      16 

Shape   .    .    . '.  =  .'...  18 

Movements      19 

Structure 19 

Undulating  Membrane 20 

Nucleus 21 

Small  Spirochaetes ...*..-  25 

Movements 25 

Structure     .....    26 

VARIABILITY  OF  FORM 29 

HABITAT 31 

CULTIVATION 31 

MULTIPLICATION  AND  DEVELOPMENT 33 

Conjugation 37 

Encystment .    ,    .   - 38 

Sporulation , 40 

Coccoid  Forms 40 

ASSOCIATION  WITH  OTHER  ORGANISMS 42 

PATHOGENICITY 45 

Relapsing  Fever . 45 

African  Tick  Fever 46 

Other  Forms  of  Current  Fevers 47 

Syphilis 48 

Yaws  (Pian,  Frambcesia)       57 

Lymphadenoma 57 

Ulcerative  Granuloma       58 

Vincent's  Angina 58 

Other  Diseases  in  Man      59 

Diseases  of  Animals  (Fowls,  Cattle,  Bat,  Pig,  Mouse)    .    .  60 

SUMMARY  OF  PHENOMENA  OF  PATHOGENICITY  OF  SPIROCH^TES  62 


8  CONTENTS. 

PAGE 

METHODS  OF  STAINING 63 

Giemsa's  Stain 63 

Other  Methods 64 

Impregnation  with  Silver 65 

Indian  Ink  Method  66 

EXAMINATION  OF  LIVING  SPIROCH^TES 66 

Dark-field  Examination 66 

Intra-vitam  Staining 67 

PSEUDO-SPIROCH^TES 67 

CLASSIFICATION ' 69 


SECTION  II. 

SYSTEMATIC  DESCRIPTION  OF  SPECIES. 

LARGE  SPIROCH^ETES     .    , 73 

Spirochaeta  Plicatilis 73 

Spirochaeta  Daxensis 76 

Spirochaeta  Balbianii 77 

Spirochaeta  Anodontae  (Spirochaeta  Spiculifera)    ....  79 

Spirochaetes  in  Various  Molluscs 82 

Spirochaeta  Pinnae •  84 

Spirochaeta  Mactrae 84 

Spirochaeta  Pectinis 86 

Spirochaeta  Interrogationis 86 

Spirochaeta  Polyspira 86 

SMALL  SPIROCH^TES  OR  SPIRILLA 86 

Spirochaeta  Obermeieri       86 

Spirochasta    Duttoni     (Spirochaeta    Novyi,     Spirochaeta 

Carteri) 88 

Spirochaeta  Pitheci 90 

Spirochaeta  Anserina      90 

Spirochaeta  Gallinarum 91 

Spirochaeta  Lagopodis 93 

Spirochaeta  Lovati .  93 

Spirochaeta  Vesperuginis 93 

Spirochaeta  Lutrae 94 

Spirochaeta  Theileri  and  Allied  Forms 94 

Spirochaeta  Suis 95 

Spirochaeta  Laverani .  96 

Spirochaeta  Muris 96 

Spirochaeta  Minor 97 

Spirochaetes  in  Mouse-cancer 97 

Spirochaeta  Gondii 98 

Spirochaeta  Culicis      98 


CONTENTS.  9 

PAGE 

Spirochaeta  Minei 98 

Spirochasta  Gadi 98 

Spirochaeta  Pelamydis 99 

Spirochaeta  Jonesii 99 

Spirochaeta  Hartmanni 99 

Spirochaeta  Bufonis ,    .    .    .    .  99 

Spirochasta  Buccalis 100 

Spirochaeta  Dentium 101 

Spirochaeta  Vincenti 102 

Spirochaeta  Gracilis -..'.' 102 

Spirochasta  Schaudinni      103 

Spirochaetes  in  Gangrenous  Processes,  Haemoptysis,  etc.  .  103 

Spirochaeta  Pseudopallida 104 

Spirochaeta  Bronchialis 104 

Spirochaeta  Microgyrata    ....... 105 

Spirochaetes  in  Small-pox      ..............  105 

Spirochaetes  in  Vaccine  Lymph    .......  ' .    .        .106 

Spirochaeta  Refringens 106 

Spirochasta  Balanitidis 107 

Spirochaeta  Pallida ...'...  108 

Spirochaeta  Pertenuis 112 

Spirochaeta  Obtusa 113 

Spirochaeta  Lymphatica 113 

Spirochaeta  Aboriginalis 114 

Spirochaeta  Interrogans 114 

VARIOUS  DOUBTFUL  SPIROCHAETES  AND  SPIRILLA 115 

BIBLIOGRAPHY 117 

BIBLIOGRAPHIC  INDEX 137 

INDEX 143 


SPIROCH/ETES. 

A  REVIEW  OF  RECENT  WORK. 

SECTION   I. 

GENERAL  CHARACTERS  OF  SPIROCH^TES. 
BIOLOGICAL  POSITION  OF  SPIROCH/ETES. 

The  name  Spirochaeta  was  first  invented  by  Ehren- 
berg  in  1833  for  the  organism  discovered  by  him  and 
named  Spirochceta  plicatilis.  It  was  regarded  by  him 
as  a  protozoon,  but  its  relationship  to  other  members  of 
this  order  was  not  clearly  determined;  and  while  some 
writers  considered  that  spirochaetes  were  members  of 
the  class  Trypanosomidae,  others  regarded  them  as 
really  bacteria.  Indeed,  in  some  of  the  earlier  text- 
books of  bacteriology  the  term  spirochaete  was  applied 
to  a  number  of  spirillar  forms. 

The  great  increase  of  interest  which  has  recently  been 
aroused  in  the  spirochaetes  dates  from  the  discovery  by 
Schaudinn  in  1905  of  the  spiral  organism  associated 
with  syphilis,  which  he  assigned  to  this  class.  The 
attention  thus  directed  to  these  minute  organisms  has 
resulted  in  the  discovery  of  a  large  number  of  forms, 
some  associated  with  conditions  of  disease,  others  living 
apparently  as  harmless  saprophytes.  Among  the  latter 
must  be  classed  the  large  varieties  present  in  several 
genera  of  shell-fish. 

The  relationship  existing  between  the  larger  and  the 
smaller  spirochaetes  has  not  been  thoroughly  eluci- 


12  SPIROCILETES. 

dated.  Indeed,  owing  to  the  minute  size  and  feeble 
staining  properties  of  the  latter  organisms,  it  is  difficult 
to  recognise  more  than  their  shape  and  movements. 
Details  of  finer  structure,  which  might  throw  light  on 
their  true  nature,  are  as  yet  wanting.  The  type  of  the 
smaller  spirochaetes  may  be  seen  in  the  organism  associ- 
ated with  the  disease  called  relapsing  fever,  usually 
called  Spirillum  or  Spiroch&ta  obermeieri,  after  its  dis- 
coverer, but  perhaps  more  correctly  on  the  basis  of 
recent  biological  nomenclature,  Spirochceta  recurrentis, 
the  spirochaete  of  recurrent  fever.  This  organism  was 
for  many  years  regarded  as  a  bacterium  and  placed  in 
the  same  class  as  the  spirillum  or  vibrio  of  cholera.  It 
does  not  seem  proved  so  far  that  this  classification  is 
incorrect,  and  the  question  is  still  at  issue,  names  of 
weight  being  found  on  both  sides  in  the  controversy 
as  to  the  protozoan  or  bacterial  nature  of  the  smaller 
spirochaetes.  Even  the  larger  spirochaetes  have  by  some 
been  assigned  to  the  bacteria  as,  for  example,  by 
Schwellengrebel,  who  points  out  the  close  resemblance 
between  an  organism,  such  as  Sp.  balbianii  and  some  of 
the  larger  spirilla,  such  as  Spirillum  giganteum  (Spm. 
volutans) .  I  hope  to  show  in  subsequent  pages  reasons 
for  thinking  that  this  view  is  probably  correct. 

With  regard  to  the  larger  spirochaetes,  it  is  certainly 
natural  to  anyone  who  observes  the  extreme  activity 
of  movement  exhibited  by  such  an  organism  as  Sp. 
anodontcz,  to  regard  it  as  prima  facie  an  animal.  This 
vigour  of  movement,  however,  does  not  by  itself  con- 
stitute an  argument  of  appreciable  weight,  for  among 
the  spirilla,  which  are  generally  admitted  to  be  bacterial, 
very  active  movement  may  be  observed.  Nevertheless, 
there  is  some  difference  between  the  two  classes  in  this 
respect;  for  while  the  spirochaetes  perform  vigorous 
lashing  movements  in  which  the  whole  body  is  bent  to- 


GENERAL   CHARACTERS   OF   SPIROCTLETES.  13 

and-fro  like  the  lash  of  a  whip,  the  spirilla  confine  them- 
selves almost  entirely  to  a  rapid  "cork-screw"  motion, 
with  little  alteration  in  the  long  axis  of  the  body.  A 
certain  degree  of  the  latter  motion  may,  however,  at 
times  be  observed  in  them,  so  that  this  difference  is 
rather  quantitative  than  one  of  kind. 

Failure  to  cultivate  spirochaetes,  large  or  small,  in 
artificial  media  has  been  urged  as  a  reason  for  regard- 
ing them  as  protozoa  and  not  bacteria,  but  this  argu- 
ment again  is  of  little  weight,  since,  not  all  known 
bacteria  can  be  grown  artificially  (e.g.,  B.  leprcs),  and 
the  quality  of  adaptability  to  cultivation  in  the  labora- 
tory can  hardly  be  looked  upon  as  essential  to  this  or 
any  other  class.  Indeed,  some  of  the  protozoa  can  be 
cultivated;  for  example,  some  forms  of  amoebae  and  of 
flagellata. 

The  curious  arrangement  of  the  staining  material 
in  the  body  of  the  larger  spirochaetes,  which  has  been 
regarded  as  constituting  a  special  nucleus,  has  been  held 
to  prove  their  animal  nature ;  but  there  is  at  present  too 
much  uncertainty  as  to  the  constitution  of  this  staining 
substance  and  its  formation  in  these  organisms  to  allow 
of  any  great  weight  being  assigned  to  this  peculiarity 
for  purposes  of  classification.  Schwellengrebel  states 
that  a  spiral  filament  similar  to  that  found  in  Sp.  bal- 
bianii  may  be  found  in  Spirillum  giganteum,  though  his 
statement  has  not  so  far  been  confirmed.  Certainly  a 
resemblance  exists  between  the  form  taken  by  the 
staining  substance  in  this  spirillum  and  that  seen  in 
some  spirochaetes  (Figs.  8,9,10). 

Other  peculiarities  of  the  large  spirochaetes  which  are 
regarded  as  protozoan  characteristics  are  the  asserted 
existence  of  an  undulating  membrane,  the  lack  of 
flagella,  and  the  habit  of  longitudinal  division  for  pur- 
poses of  multiplication.  With  regard  to  the  first  of 


14  SPIROCILETES. 

these,  doubt  has  recently  been  thrown  on  the  identity 
of  the  appearance  seen  in  spirochaetes  with  the  true 
undulating  membrane  of  such  organisms  as  trypano- 
somes.  Consequently  it  is  premature  to  regard  this  as 
a  distinctive  feature.  Flagella  have  been  described  in 
some  of  the  smaller  spirochaetes,  but  with  doubtful 
certitude ;  they  appear  to  be  absent  in  the  larger  forms. 
Such  absence  does  not  seem  to  constitute  an  argument 
for  or  against  their  vegetable  nature.  The  method  of 
reproduction  even  in  the  larger  spirochaetes  is  still  dis- 
puted, but  there  appears  to  be  a  balance  of  authority 
in  favour  of  longitudinal  division  as  the  usual  occurence, 
and  I  have  myself  seen  an  instance  of  what  appeared  to 
be  this  procedure  in  Sp.  anodonta. l  The  occurrence  of 
encysted  forms  in  such  species  as  Sp.  balbianii  has  also 
been  held  to  prove  them  to  be  protozoa,  but  the  real 
nature  of  the  curled  up  forms  is  as  yet  uncertain,  and 
it  is  not  unknown  for  vegetable  organisms  to  enter  upon 
a  resting  stage. 

The  strongest  argument  which  could  be  brought  for- 
ward in  favour  of  the  animal  nature  of  the  spirochaetes 
would  be  the  occurrence  of  conjugation — a  distinctive 
protozoan  characteristic.  This  phenomenon  has  been 
described  as  occurring  in  several  species  of  spirochaetes 
(see  p.  37),  but  the  evidence  adduced  is  unconvincing. 

The  nature  of  the  smaller  spirochaetes  is  still  more  un- 
certain. The  points  about  them  which  seem  to  place 
them  in  the  same  class  as  the  larger  spirochaetes  are 
their  general  shape  and  movements,  the  occurrence  of 
longitudinal  division,  the  asserted  possession  by  some  of 
them  (not  by  all)  of  an  undulating  membrane,  and  the 
absence  of  flagella.  Their  resemblance  to  the  larger 
forms  in  the  matter  of  general  shape  and  movements  is 

1  If  Schmeidlechner's  observation  of  bacilli  which  divide  lon- 
gitudinally is  confirmed,  this  mode  of  multiplication  cannot  be 
considered  a  distinguishing  feature  of  animal  organisms. 


GENERAL  CHARACTERS   OF  SPIROCILETES.  15 

certainly  striking,  but  is  not  in  itself  a  strong  argument 
for  combining  them  in  a  single  group.  The  possession 
of  an  undulating  membrane  by  these  minute  organisms 
is  certainly  doubtful,  and  the  presence  or  absence  of 
flagella  has  little  weight  in  this  connection,  except  as  an 
arbitrary  ground  of  classification.  The  question  as  to 
the  multiplication  of  these  little  organisms  by  lon- 
gitudinal or  transverse  division  is  still  undecided;  but 
the  formation  of  a  fine  thread  at  the  point  of  divi- 
sion seems  to  be  established,  whereas  bacteria  divide  by 
means  of  a  distinct  septum.  The  argument  from  the 
possibility  of  cultivation  on  artificial  media  has  just 
been  noticed  to  be  valueless,  and  indeed  some  of  the 
small  spirochaetes  have,  if  the  claims  of  various  writers 
are  to  be  believed,  been  successfully  grown  in  the  lab- 
oratory. The  conveyance  of  some  of  the  pathogenic 
spirochaetes  by  biting  insects  (ticks)  has  been  used  as 
a  proof  of  their  animal  nature,  but  can  hardly  be  re- 
garded as  a  conclusive  argument ;  for  it  has  not  been 
proved  that  they  undergo  any  constant  process  of 
development  within  these  hosts. 

A  stronger  argument  has  been  drawn  from  the  action 
of  certain  chemical  reagents  on  small  spirochaetes.  Thus 
it  is  said  that  cobra-venom,  taurocholate  of  sodium, 
saponin,  and  pancreatic  extract  destroy  spirochaetes 
and  protozoa,  but  do  not  affect  bacteria.  Spirochaetes 
are  little  affected,  however,  by  distilled  water  or  by 
concentrated  salt-solution.  It  is  also  asserted  that 
whereas  bacteriolysins  are  thermostable  (not  destroyed 
by  heat),  the  corresponding  substances  which  act  on 
spirilla  (spirochsetes)  are  thermolabile,  thus  resembling 
the  haemolysins.  On  the  other  hand,  the  action  of 
immune  serum  in  producing  agglomeration  or  aggluti- 
nation of  spirochaetes  is  like  that  exerted  upon  bacterial 
organisms;  and  Sp.  pallida — if  this  be  really  the  cause 


1 6  SPIROCILETES. 

of  syphilis — gives  rise  in  the  body  of  the  human  host  to 
the  formation  of  an  antibody  (copula  or  amboceptor)  as 
do  the  pathogenic  bacteria.  This  form  of  immune  sub- 
stance has  not  been  found  in  infections  caused  by 
protozoa. 

It  is  to  be  hoped  that  further  study  of  the  life-history 
and  development  of  spirochsetes  will  throw  definite 
light  on  their  true  biological  position.  If  the  observa- 
tjon  of  Leishman  on  the  development  of  Sp.  duttoni 
into  coccoid  forms  be  confirmed,  and  if  a  similar  change 
can  be  shown  to  take  place  in  other  spirochaetes,  as  is 
suggested  by  my  own  experience,  their  close  relationship 
to  the  bacteria  will  be  manifest.1 

In  attempting  to  assign  the  spirochastes  either  to 
the  protozoa  or  the  bacteria,  it  must  be  borne  in 
mind  that  there  are  several  other  kinds  of  organisms 
known,  about  which  there  is  equal  uncertainty  as  to 
their  correct  classification.  Indeed,  it  has  to  be  con- 
fessed that  there  are  no  definite  criteria,  by  which  to 
draw  a  line  of  division  between  the  lowest  forms  of 
animals  and  plants.  This  must  be  regarded  as  al- 
most inevitable,  since  both  animal  and  vegetable 
kingdoms  present  to  us  as  their  lowest  members  uni- 
cellular organisms  of  very  simple  structure,  while  it 
is  at  least  probable  that  at  the  beginning  of  things  a 
single  form  of  living  matter  gave  origin  to  both  the 
great  divisions,  animals  and  plants. 

MORPHOLOGY  OF  SPIROCHAETES. 

The  minute  structure  of  the  larger  spirochaetes  has 
been  carefully  observed  in  the  species  Sp.  balbianii  and 

1  The  suggestion  that  spirochaetes  are  stages  in  the  development 
of  a  bacterial  organism  is  referred  to  on  page  42.  Schaudinn's 
original  view  that  they  constitute  a  stage  in  the  life-history  of  a 
trypanosome  is  now  generally  supposed  to  have  been  based  on  a 
mistaken  identification.  (See  Novy  and  MacNeal,  Journ.  Infect. 
Dis.,  1905,11,256;  Sergent,  Edm.  and  M.,  Ann.  d.  VInst.  Pasteur, 
1907.) 


GENERAL   CHARACTERS    OF   SPIROOELETES.  17 

Sp.  anodonta,  which  are  easily  obtainable,  as  well  as,  to 
a  less  extent,  in  the  typical  species  Sp.  plicatilis.     De- 


FIG.  i.  FIG.  2.  FIG.  3. 

FIG.  i. — Spirochaeta  plicatilis :     Schematic.      (After  Doflein.) 
PIG.   2. — Spirochaeta  balbianii,   showing  so-called  "undulating 

membrane."      (Perrin.) 

FIG.  3. — Spirochaeta  anodontae,  with  "undulating  membrane." 

(Keysselitz.) 

scribed  in  general  terms,  they  are  elongated,  vermi- 
form organisms,  endowed  with  active  movement.  They 
are  said  to  be  flattened  from  above  downward,  thus 


l8  SPIROCILETES. 

resembling  strips  of  ribbon  rather  than  threads  of  cot- 
ton in  form.  They  are  further  stated  to  be  furnished 
with  an  undulating  membrane  and  with  a  complicated 
nuclear  apparatus  (see  Figs,  i,  2,  3). 

Shape. — The  length  of  these  organisms  is  usually 
not  less  than  twenty  times  the  breadth,  Sp.  balbianii 
for  example  reaching  a  length  of  TOO/*,  while  its  breadth 
is  usually  less  than  5/1.  The  length  of  Sp.  plicatilis  may 
reach  500^.  I  have  found  specimens  of  Sp.  anodontcs 
nearly  130;*  in  length,  without  any  signs  of  division  at 
any  point ;  but  unusually  long  forms  are  regarded  by 
some  writers  as  about  to  enter  upon  the  process  of  fission. 
It  is  difficult  to  ascertain  whether  these  organisms 
are  naturally  flattened  in  form,  as  during  active  move- 
ment, while  they  are  alive,  the  exact  contour  cannot  be 
made  out,  while  some  flattening  might  easily  take  place 
after  death  in  the  process  of  fixing.  Sp.  balbianii  is 
said  by  Perrin  to  be  round  in  section. 
Fantham  and  Gross  have  succeeded  in 
obtaining  transverse  sections  of  spiro- 
Cn3etes>  which  suggest  a  form  such  as  that 
chaete  in  trans-  shown  diagrammatically  in  figure  4 — a 

verse  section.  ..    ,  .  .  ,  ..  ,         , 

rounded  organism  with  a  loose  sheath 
which  may  project  on  one  side.  The  organism  might 
thus  appear  flattened  in  one  direction,  and  round  in 
another;  and  the  opposing  views  might  thus  be  har- 
monised. Gross  regards  the  projecting  portion  as  con- 
stituting a  crest  or  comb. 

The  ends  of  the  organism  may  be  either  blunt  or 
pointed  (Figs.  2,  3).  It  is  uncertain  whether  this  fea- 
ture is  always  constant  in  the  same  species.  Thus,  in 
the  case  of  Sp.  anodonta  Schellack  would  constitute  the 
individuals  with  pointed  ends1  a  separate  species  (Sp. 

*Cf.  Castellani's  distinction  of  Sp.obtusa  and  Sp.  acuminata, 
page  113. 


GENERAL   CHARACTERS    OF   SPIROCILETES.  IQ 

spiculifera) .  Certainly  this  form  tends  to  be  shorter 
on  the  average  than  the  form  with  blunt  extremities; 
but  some  overlapping  may  occur,  and  it  seems  prema- 
ture to  make  a  specific  division  on  this  ground  alone 
(see  page  81). 

Movements. — The  movements  of  the  large  spirochaetes 
are  very  rapid  under  normal  conditions,  but  become 
slower  as  vitality  diminishes.  They  are  of  three  main 
kinds:  (i).  Lashing  movements,  such  as  are  performed 
by  an  eel  suspended  on  a  hook,  the  whole  body  being 
bent  from  side  to  side,  taking  the  forms  of  a  circle  or  an 
S  or  a  figure  of  eight.  These  are  most  evident  when  the 
spirochaete  is  attached  in  some  way  by  one  extremity  or 
hindered  in  its  progression.  (2).  Undulating  move- 
ments, which  are  compared  to  the  flapping  of  a  sail 
"in  the  wind,"  a  wave  passing  from  one  end  of  the 
organism  to  the  other  and  being  rapidly  succeeded  by 
other  similar  waves  in  the  same  direction.  It  would 
seem  that  the  direction  may  be  reversed.  (3).  A  rota- 
tory movement,  like  that  of  a  cork-screw  when  pushed 
into  a  cork,  the  whole  spirochaete  turning  on  a  longit- 
udinal axis  passing  through  the  centres  of  the  spirals 
(the  axis  of  the  helix).  Special  forms  of  movement 
are  described  by  Perrin  as  taking  place  in  the  process  of 
encystment,  one  end  of  the  organism  gliding  down  the 
side  of  the  body,  to  be  followed  by  a  similar  movement 
of  the  other  extremity,  and  finally  by  a  continuous 
movement  by  which  the  creature  is  rolled  upon  itself. 

Structure. — Examined  microscopically,  the  structure 
of  the  spirochaete  is  seen  to  consist  of  an  external  coat 
or  periplast,  which  contains  an  inner  protoplasmic  sub- 
stance or  entoplasm.  The  periplast  may  be  prolonged 
at  one  or  both  ends  of  the  organism  into  an  appendage, 
which  some  authors  regard  as  a  flagellum.  This  iden- 
tification seems  incorrect.  It  has  also  been  suggested 


20  SPIROCILETES. 

that  these  appendages  are  formed  in  the  process  of 
division  of  the  spirochaste,  and  thus  may  not  be  per- 
manent. Perrin  speaks  of  the  periplast  retaining  its 
form  after  the  entoplasm  has  escaped  from  within  it 
and  become 'encysted. 

Attached  to  the  periplast  or  forming  part  of  it  is  the 
so-called  undulating  membrane  (Figs.  2,  3).  This  is 
described  as  running  round  the  body  of  the  spirochaete 
in  a  spiral  direction,  and  as  being  bounded  at  its  external 
margin  by  a  darkly  staining  fibril  (randfibrille) ,  while 
other  fibrils  may  be  seen  running  in  it  parallel  to  the 
marginal  one.  The  latter  is  said  to  be  connected  at  its 
extremities  with  two  darkly  staining  particles,  situated 
at  the  ends  of  the  spirochaete,  called  by  Gonder  bleph- 
aroplasts,  on  the  analogy  of  the  blepharoplast  of 
trypanosomes. 

The  nature  of  this  undulating  membrane  has  been 
called  in  question.  Laveran  and  Mesnil  regard  the 
appearance  thus  named  as  produced  by  the  existence 
of  a  loose  sheath  outside  the  spirochaste ;  and  a  sheath 
has  been  described  in  several  of  the  smaller  spiro- 
chaetes  by  different  writers  (see  p.  26).  Schellack 
believes  that  the  so-called  undulating  membrane  is  an 
artifact,  produced  in  the  process  of  fixation.  My  own  ex- 
perience with  Sp.  anodontcs  would  seem  to  support  this 
view,  as  I  find  that  the  better  the  specimen  is  fixed,  the 
less  frequent  are  individuals  which  appear  to  have 
undulating  membranes.  Nor  does  the  appearance 
when  present  correspond  at  all  closely  with  that  seen 
in  trypanosomes,  the  marginal  fibril  being  often  seen 
as  a  separate  structure  with  no  membrane  visible 
between  it  and  the  body  of  the  spirochaete,  and  often 
also  appearing  at  only  one  or  two  points,  not  as  a  struc- 
ture continuous  all  along  the  organism.  The  appear- 


GENERAL   CHARACTERS    OF    SPIROCILETES.  21 

ance  of  this  fibrillary  structure  at  the  outer  margins  of 
the  curls  presented  by  the  spirochaste  suggests  that  it 
may  be  a  portion  of  periplast  which  has  become  fixed 
to  the  slide,  and  has  thus  remained  fast,  in  a  figure  cor- 
responding with  the  curves  in  which  the  creature  lay  at 
the  moment  of  death:  then  as  drying  took  place  and 
the  body  of  the  spirochaete  shrank,  the  latter  straight- 
ened out  somewhat,  while  the  adherent  periplast  re- 
mained. Sometimes,  however,  the  "membrane"  runs 
in  a  straight  line  between  neighbouring  coils,  having 
apparently  shrunk  more  than  the  body-substance. 

Borrel  and  Cernovodeanu,  however,  state  that 
they  have  seen  an  undulating  membrane  in  living 
specimens  of  Sp.  anodontcB. 

Perrin  thinks  that  the  undulating  membrane  is  not 
of  much  importance  as  an  aid  to  locomotion  in  Sp. 
balbianii. 

With  regard  to  the  structure  of  the  entoplasm  and  its 
contents  an  equal  degree  of  uncertainty  prevails. 
Perrin  described  in  Sp.  balbianii  the  existence  of  pecu- 
liar form  of  nucleus.  It  consists,  according  to  this  writer 
of  a  spiral  band  of  achromatic  substance  (karyosome) 
on  which  appear  at  intervals  masses  of  staining  material 
(chromatin  ?) .  These  latter  usually,  in  resting  organ- 
isms, take  the  form  of  rods  arranged  so  as  to  lie  trans- 
versely, more  or  less  at  right  angles  to  the  main  axis  of 
the  spirochaete  (Fig.  5) ;  but  when  fission  is  about  to 
occur,  the  rods  assume  other  forms  and  undergo  divi- 
sion. Fantham  confirmed  Perrin's  statements.  Keys- 
selitz  described  in  Sp.  anodonta  balls  and  rods  of  chro- 
matic material  (Fig.  6) ,  and  thinks  that  there  may  be  a 
spiral  arrangement  such  as  Perrin  found,  but  did  not 
himself  observe  it.  Schellack  confirms  the  existence  of 
bands  of  chromatin,  but  denies  that  they  are  connected 


22  SPIROCILETES. 

by  any  spiral  thread.  He  regards  the  spirochaete  as 
made  up  of  a  series  of  chambers,  in  which  these  rod- 
shaped  nuclei  lie,  and  thinks  that  the  increase  in  length 
of  the  spirochaete  is  effected  by  a  multiplication  of  these 


FIG.  5.  FIG.  6. 

FIG.  5. — Spirochaeta  balbianii,  showing  nuclear  (?)  bands. 
(Perrin.) 

FIG.  6. — Spirochseta  anodontse,  with  irregular  masses  of  staining 
substance.  (Keysselitz.) 

chambers,  taking  place  especially  at  the  ends.  In 
some  cases  the  staining  substance  appears  as  a  central 
continuous  band  or  rod,  running  along  the  middle  of 
the  body  of  the  spirochaete  (Fig.  7). 


GENERAL   CHARACTERS   OF   SPIROCILETES.  23 

The  appearances  seen  in  Sp.  plicatilis  are  some- 
what different  (Fig.  i).  Here  the  organism  appears  to 
be  furnished  with  a  series  of  rounded  nuclei,  distributed 


FIG.  7. — Spirochseta  balbianii;   rod-shaped  "nucleus."     (Perrin.) 

fairly  evenly  throughout  its  length.  The  general  out- 
line of  the  spirochaete  is  also  different,  since  it  shows 
not  only  the  large  undulations  seen  in  Sp.  anodonta 
and  balbianii,  but  also  a  series  of  smaller  waves  super- 
imposed upon  the  others.  It  has  no  undulating  mem- 


FIG.  8. — Spirochaeta    anodontae,    showing   irregular    granules    of 
staining  material. 

brane,  as  depicted  in  Doflein's  illustration — indeed, 
judged  merely  by  a  comparison  of  this  with  the  illus- 
trations of  Sp.  balbianii,  given  by  Perrin  or  Fantham,  it 


SPIROCILETES. 


is  difficult  to  believe  that  the  two  organisms  belong  to 
the  same  genus. 

So  far  as  I  can  judge  from  my  own  observations,  the 
peculiar  aggregations  of  staining  material  seen  in  Sp. 


FIG.  9.  FIG.  10. 

FIG.  9. — Bacteria  from  rectum  of  ox  for  comparison  with  Fig.  8. 
FIG.   10. — Spirillum  volutans  for  comparison  with  Fig.   8. 

anodonta  are  not  to  be  looked  on  as  nuclei.  They  are 
irregular  in  shape  and  quite  unlike  the  nuclei  of  pro- 
tozoa. They  seem  to  occur  for  the  most  part  in  organ- 
isms which  are  degenerating  (so-called  "involution 


9 


FIG.  ii. — Filaments  of  algae:  a,  Resembling  fusiform  bacilli;  b, 
showing  central  staining  rod;  c,  with  series  of  chambers;  d,  e, 
curved  like  spirochaetes. 

forms  ") .  In  other  instances  they  may  point  to  a  forth- 
coming developmental  change  (p.  40).  The  great 
majority  of  the  organisms  in  my  experience  stain 


GENERAL   CHARACTERS    OF    SPIROCH^TES.  25 

homogeneously  like  bacteria.  In  some  a  vacuolated 
structure  may  be  made  out,  resembling  that  seen  in 
some  of  the  larger  spirilla.  In  the  larger  bacteria  also 
granules  or  masses  of  staining  material  may  be  seen,  not 
unlike  those  in  Sp.  anodontce  (see  Figs.  8,  9,  10).  Fila- 
ments of  algae  may  also  show  features  bearing  a  close 
resemblance  to  some  of  those  described  in  spirochaetes 
(Fig.  n). 

In  the  case  of  the  smaller  spirochaetes  the  minute 
size  of  the  organisms  renders  the  investigation  of  their 
intimate  structure  almost  impossible.  The  general 
shape  is  the  same  as  that  of  the  large  varieties,  accord- 
ing to  most  writers,  but  it  is  even  more  difficult  to 
make  sure  of  their  flattened  shape  than  in  the  larger 
organisms.  Some  are  definitely  stated  to  be  round  in 
section,  e.g.,  Sp.  pallida,  but  many  authors  follow 
Vuillemin  and  Schaudinn  in  placing  this  and  a  few 
others  in  a  separate  genus,  Treponema,  partly  on  this 
account. 

The  movements  of  the  small  spirochaetes  are  the 
same  as  those  described  above.  An  additional  form 
of  movement  is,  however,  described  by  Plaut  in  Sp. 
vincenti  and  called  by  him  "euglenoid"  movement: 
it  consists  in  the  appearance  of  a  thickening  at  one 
point  in  the  body  of  the  organism  and  the  gradual  pas- 
sage of  this  swelling  from  one  end  to  the  other.  Krzys- 
talowicz  and  Siedlecki  describe  a  somewhat  similar  com- 
pression and  extension  of  the  body  of  Sp.  pallida  when  it 
is  in  motion.  It  must  be  remembered  that  in  these  mi- 
nute thread-like  organisms  it  is  impossible  to  distinguish 
under  the  microscope  between  rotatory  movement  and 
undulation.  Lashing  movements  are  said  to  be  less 
marked  in  treponema:  indeed,  the  movements  Sp. 
pallida  (Trep.  pallidum)  are  altogether  sluggish. 


26  SPIROCHjETES. 

Of  the  actual  structure  of  these  tiny  organisms  little 
can  be  said.  An  undulating  membrane  has  been  de- 
scribed in  some  species,  as  by  Gonder  in  Sp.  vesperu- 
ginis,  by  Prowazek  in  Sp.  gallinarum,  by  Hoffmann  in 
Sp.  buccalis,  and  by  Schaudinn  in  Sp.  refringens.  On 
the  other  hand,  Levaditi  denies  that  Sp.  refringens 
possesses  an  undulating  membrane,  and  this  structure 
has  never  been  seen  or  described  in  Sp.  obermeieri, 
Sp.  pallida,  Sp.  muris,  the  spirochaete  of  ulcerating 
cancer,  etc.  When  it  is  borne  in  mind  that  one  of 
the  favourite  stains  for  spirochaetes,  Giemsa's, 
is  deposited  round,  as  well  as  in,  the  struct- 
ures which  attract  it,  it  is  easy  to  imagine  a 


FIG.  13.  FIG.  14. 

FIG.  12.— Spirochaetaduttoni;  appearance  of  a  sheath.  (Stephens.) 
FIG.   13. — Spirochaeta  buccalis.    Sheath.  X22SO.     (Prowazek.) 
FIG.  14.— Anthrax  bacillus,  with  appearance  of  a  sheath.  (Ascoli.) 

possible  source  of  error  in  observing  this  point  in  a 
minute  organism  of  spiral  shape.  Most  of  the  illustra- 
tions of  undulating  membranes  in  these  small  spiro- 
chaetes are  far  from  convincing. 

A  definite  sheath  has  been  described  in  some  forms, 
e.g.,  in  Sp.  duttoni  by  Stephens  (Fig.  12),  in  Sp.  pallida 
by  Leuriaux  and  v.  Geets,  in  Sp.  buccalis  by  Prowazek 
(Fig.  13),  in  the  spirochaste  found  by  Baruchello  and 
Pricolo  in  equine  pneumonia,  and  in  an  unidentified 
spirochaete  found  by  Kenrick  in  a  case  of  fever.  An 
appearance  resembling  a  sheath,  may  be  found  in  bac- 
teria, as  is  shown  in  the  accompanying  illustration  of 


GENERAL   CHARACTERS    OF   SPIROCH^TES.  27 

Bacillus  anthracis  (Fig.   14),  taken  from  a  paper  by 
Ascoli.1 

Terminal  flagella  are  described  in  some  forms,  e.g., 
Sp.  obermeieri  (Reichert),  and  Sp.  duttoni  (Stephens), 


FIG..  15. — Spirochaeta   pallida,  with  terminal  filament;   so-called 
autogamy  stage.      (Prowazek.) 

but  these  are  generally  regarded  (as  by  Schaudinn  and 
by  Krzystalowicz  and  Siedlecki)  as  mere  prolongations 
of  the  periplast  (Figs.  15,  16,  17,  18).  Lateral  flagella 
have  also  been  described  (e.g.,  by  Fraenkel  in  Sp. 


FIG.  16.  FIG.  17.         FIG.  18.  FIG.  19. 

FIG.   1 6. — Spirochaeta,  media  of  stomatitis,  with  terminal  fila- 
ment.    X225O.     (Prowazek.) 

FIG.  17.— Spirochseta  duttoni  (terminal  filament).     (Schellack.) 

FIG.    18. — Spirochaeta     balbianii,     with     terminal     "brush." 
(Schellack.) 

FIG.   19. — Spirochaeta       recurrentis;     splitting     of      penplast. 
(Schellack.) 

obermeieri},  but  are  probably  due  to  injury  in  fixation 
of  specimens  (Fig.  19). 

Small  granules  or  nodules,  appearing  rather  larger  in 
diameter  than  the  rest  of  the  spirochaete,  are  commonly 

*Centralbl.f.  Bakt.,  1908,  XLVI,  186. 


28  SPIROCH^TES. 

seen.  A  single  terminal  nodule  has  been  noted,  for 
example,  in  Sp.  duttoni  (Button,  Todd  and  Tobey),  Sp. 
pallida  (Dudgeon),  Sp.  pertenuis  (Castellani) ,  and  in 


FIG.  20.  FIG.  21. 

FIG.  20.  —  Spirochaete  of  ulcerating  cancer  (left)  and  Sp.  refringens 
(right),  with  terminal  nodules.  (Schaudinn.) 

PIG.  21.  —  Spirochaeta  pertenuis  with  terminal  granules. 
(Castellani.) 

many  other  forms  (Figs.  20,  21,  22,  23).  Two  nodules 
may  be  placed  toward  the  same  extremity  (e.g.,  Sp. 
aboriginalis,  Bosanquet)  .  One  or  more  of  such  dots  or 
masses  may  also  be  seen  (Figs.  24,  25),  in  the  central 


FIG.  22.      FIG.  23.  FIG.  24.  FIG.  25. 

FIG.  22.  —  Spirochasta  aboriginalis  with  granules  and  "loop." 
(Bosanquet.) 

FIG.  23.  —  Spirochaeta  duttoni.     Injured  specimen.      (Schellack.) 

FIG.  24.  —  Spirochaeta  duttoni,  with  masses  of  staining  sub- 
stance. Supposed  to  represent  nuclei.  X22oo.  (Mayer.) 

FIG.  25.  —  Spirochaeta  lagopodis,  showing  dots  resembling  nuclei. 
(Fantham.) 

part  of  the  organism.  These  have  been  supposed  to 
represent  nuclei,  and  Herxheimer  has  indentified  a 
kineto-nucleus,  a  tropho-nucleus  and  centrosome  —  a 


GENERAL   CHARACTERS    OF   SPIROCILETES.  29 

degree  of  refinement  which  at  present  seems  fanciful. 
These  little  nodular  bodies  sometimes  appear  to  be 
attached  to  the  side  of  the  spirochsete  rather  than  to 
lie  in  its  body  (Fig.  26).  It  is  not  unusual  to  find 
spirochastes  which  resemble  a  chain  of  granules  set  like 
a  string  of  beads  (Fig.  27) :  these  are  usually  regarded 
as  degenerated  forms  ("moniliform  degeneration"),  but 
the  possibility  that  these  organisms  break  up  into 


FIG.  26.  FIG.  27. 

FIG.  26. — Spirochaeta  gallinarum,  with  lateral  nodule.  (Pro- 
wazek.) 

FIG.  27. — Spirochaeta  obermeieri,  showing  breaking  up  of  body 
into  rods  and  granules  (diagrammatic). 

minute  bodies  capable  of  growing  into  other  individu- 
als, when  placed  in  favourable  circumstances,  cannot 
be  excluded. 

A  loop  has  been  described  as  present  at  one  end  of 
the  spirochaete  in  certain  forms,  e.g.,  in  Sp.  pallida 
(Reuter),  Sp.  pertenuis  (Castellani) . 

Masses  of  organisms  may  be  found  tangled  together, 
as  in  Sp.  pallida  (Bandi  and  Simonelli),  Sp.  dentium 
(Miller),  Sp.  obermeieri  (Zettnow),  and  Sp.  duttoni 
(in  the  tick,  Koch) .  I  have  once  seen  a  large  number 
of  Sp.  anodontcz  forming  a  tangle. 

VARIABILITY  OF  FORM. 

A  very  important  question  is  that  of  the  possibility 
of  some  degree  of  change  of  form  in  a  single  species, 
since  upon  the  negative  reply  to  this  question  must  at 
present  depend  the  possibility  of  accurately  distin- 


30  SPIROCIOETES. 

guishing  and  classifying  these  organisms.  The  exist- 
ence of  somewhat  different  forms  side  by  side  has  been 
already  referred  to  in  connection  with  Sp.  anodontaz. 
It  is  remarkable  that  two  forms  also  exist  together  in 
Tapes  laeta  (Sp.  tapetos)  and  in  other  molluscs  (Schellack) . 
The  coexistence  of  the  two  forms,  Sp.  pallida  and  Sp. 
refringens,  in  syphilitic  lesions  is  well  known.  Two 
forms  are  described  by  Loewenthal  in  ulcerated  cancers , 
one  larger  and  the  other  smaller,  and  two  also  by 
Branch  in  cases  of  haemoptysis,  and  by  Moritz  in 
decaying  potato  (5^?.  polyspird).  Castellani  found 
four  forms  associated  in  his  cases  of  haemoptysis. 

The  question  of  change  of  form  has  been  most  care- 
fully studied  in  connection  with  S^.  pallida.  Bertarelli 
and  Volpino  describe  intermediate  forms  between  Sp. 
pallida  and  Sp.  refringens,  and  Bosc  suggests  that 
refringens  is  a  degenerative  form  of  pallida.  Schere- 
schewsky  noted  irregular  forms  of  Sp.  pallida  in  cul- 
tures, and  was  struck  by  the  resemblance  of  some  of 
these  to  refringens,  and  Sobernheim  and  Tomasoli  also 
observed  irregular  forms.  5^.  pallida  is  noted  for  the 
regularity  of  its  curls  and  is  indeed  identified  by  this 
means;  yet  Schaudinn  observed  that  almost  straight 
forms  might  occur;  Sobernheim  and  Tomasoli  found 
specimens  with  straight  portions  toward  the  centre ;  and 
Fouquet  recorded  the  discovery  of  straight  forms  in 
tertiary  syphilis.  Krzystalowicz  and  Siedlecki  noted 
changes  in  Sp.  pallida  by  which  individuals  contracted 
and  thus  became  shorter  and  plumper.  Eitner,  who 
observed  living  forms  indistinguishable  from  Sp.  pallida 
noted  that  some  of  these  straightened  out  when  dead 
and  were  thus  differentiated. 

Krienitz  studied  the  spirochaetes  present  in  the  stom- 
ach and  found  that  they  underwent  a  change  of  form 
according  to  alterations  in  their  environment.  He 


GENERAL   CHARACTERS    OF   SPIROCH^TES.  31 

consequently  held  that  morphology  does  not  constitute 
a  valid  test  of  species. 

These  observations  are  sufficient  to  indicate  that  con- 
siderable doubt  must  be  felt  as  to  the  constancy  of  form 
among  the  small  spirochaetes ;  and  when  it  is  remembered 
that  the  identification  of  such  a  form  as  Sp.  pallida  de- 
pends entirely  upon  its  morphology,  and  that  many 
forms  have  been  described  in  conditions  other  than 
syphilis  which  can  only  be  differentiated  by  experts  from 
this  spirochaete,  the  uncertainty  that  must  continue  to 
prevail  as  to  relations  of  the  small  spirochastes  to  each 
other  and  to  the  morbid  conditions  with  which  they 
are  associated,  is  undeniable. 

HABITAT. 

The  only  free-living  spirochaetes  at  present  known 
are  Sp.  plicatilis  and  Sp.  daxensis  which  are  found  in 
fresh  water.  Sp.  polyspira  is  a  saprophyte  found  in 
decaying  vegetable  matter  (potato).  The  rest  of  the 
large  spirochastes  are  saprophytes  or  commensals  in  the 
alimentary  canal  of  various  shell-fish.  Of  the  smaller 
forms  many  are  apparently  saprophytes  on  the  surface 
of  ulcerative  lesions ;  others  are  also  saprophytic  within 
the  intestines  of  various  animals.  A  few  only  are 
definitely  pathogenic,  being  capable  of  multiplying 
within  the  living  tissues  of  an  animal  host  and  there 
producing  poisonous  products. 

CULTIVATION. 

Spirochaetes  have  never  been  satisfactorily  culti- 
vated; that  is  to  say,  they  have  never  been  isolated 
in  artificial  media  and  passed  from  culture  to  culture. 
Many  authors  have  stated  that  they  have  succeeded  in 
producing  multiplication  of  spirochaetes  on  laboratory 
media.  Thus  Duval  and  Todd  kept  Sp.  duttoni  alive 


32  SPIROCILETES. 

in  a  mixture  of  yolk  of  egg  and  decoction  of  mouse- 
flesh  for  forty  days.  They  observed  the  formation  of 
long  chains  under  these  conditions.  Williams  grew 
this  organism  in  defibrinated  blood,  and  Norris,  Pappen- 
heimer  and  Flournoy  cultivated  Sp.  obermeieri  in  cit- 
rated  blood.  Lebailly  states  that  Sp.  pallida  multiplied 
when  kept  on  pieces  of  infected  liver,  and  Volpino  and 
Fontana  found  that  it  even  grew  into  pieces  of  normal 
liver  placed  in  contact  with  the  infected  tissues.  Bor- 
rel  and  Burnet  cultivated  Sp.  gallinarum  in  blood. 
Levaditi  cultivated  Sp.  pallida  and  also  Sp.  refringens 
in  collodion  sacs  placed  inside  the  peritoneal  cavities  of 
animals,  but  these  cultures  were  not  pure,  being  con- 
taminated with  great  numbers  of  bacterial  organisms. 
Beer  claims  to  have  cultivated  Sp.  pallida,  anaerobically 
like  a  bacterium  on  bouillon  mixed  with  ascitic  fluid. 

Leuriaux  and  v.  Geets  state  that  they  cultivated  Sp. 
pallida  in  a  mixture  of  broth  with  cerebrospinal  fluid 
taken  from  syphilitic  patients;  after  incubation  in  this 
medium  the  precipitate  formed  was  sown  on  pigs'  se- 
rum, and  colonies  developed.  Other  writers  state  that 
they  have  obtained  multiplication  of  these  spirochaetes 
by  incubating  portions  of  infected  organs ;  and  Levaditi, 
as  previously  mentioned,  "cultivated"  Sp.  pallida  in 
collodion  sacs  within  the  peritoneal  cavities  of  mon- 
keys, but  found  the  resulting  "cultures"  incapable  of 
inducing  infection. 

Recently  successful  attempts  to  cultivate  Sp.  pallida 
are  reported  by  Schereschewsky  and  by  Miihlens.  The 
medium  used  was  horse-serum,  which  was  first  heated 
to  58°  to  60°  C. — a  point  at  which  it  threatens  to  solidify 
— and  then  kept  at  37°  C.  for  three  days.  A  piece  of 
infected  tissue  (e.g.,  a  bit  of  a  gland  taken  from  a  patient 
suffering  from  syphilis)  is  then  inserted  deeply  into  the 
medium  and  kept  under  anaerobic  conditions  for  five 


GENERAL   CHARACTERS    OF   SPIROCILETES.  33 

days  or  so,  when  multiplication  of  the  spirochaetes  is 
found  to  have  taken  place.  An  odour  of  sulphureted 
hydrogen  is  evolved,  which  may  be  compared  with 
the  foetid  odour  produced  by  the  organisms  of  Vin- 
cent's angina.  Schereschewsky  found  that  forms 
resembling  Sp.  refringens  first  developed  and  after- 
ward characteristic  pallidce.  Inoculation  of  animals 
with  the  cultures  was  unsuccessful. 

Levaditi  and  Stanesco  also  record  successes  in  culti- 
vating Sp.  gracilis  and  Sp.  balanitidis  in  a  somewhat 
similar  way.  In  the  case  of  the  latter  organism  they 
first  inoculated  tubes  of  horse-serum  and  then  made 
sub-cultures  in  collodion  sacs  containing  this  serum, 
placed  in  the  serum  tubes.  They  noted  that  multi- 
plication of  the  spirochaetes  only  occurred  when  the 
media  were  liquefied  by  the  action  of  proteolytic 
bacteria — an  example  of  symbiosis  which  they  compare 
with  that  needed  for  the  cultivation  of  amoebae.1 

Most  recently  of  all  Bruckner  and  Galacesco  have 
grown  Sp.  pallida  on  coagulated  ascitic  fluid  and  have 
successfully  inoculated  rabbits  with  sub-cultures  made 
a  month  after  the  material  had  been  sown  on  artificial 
medium. 

MULTIPLICATION  AND  DEVELOPMENT. 

Much  controversy  has  centred  round  the  question  of 
the  mode  of  division  of  spirochaetes,  large  and  small. 
Perrin  described  longitudinal  division  in  Sp.  balbianii, 
and  Keysselitz  in  Sp.  anodontcs;  and  I  have  seen  what 
appeared  to  be  this  process  taking  place  in  the  latter. 
Schwellengrebel  believes  that  transverse  division  is  the 
rule  in  Sp.  balbianii,  and  Schellack  finds  it  in  Sp.  ano- 
dontcs (Fig.  28)  and  in  other  forms.  Among  the  smaller 

'The  "symbiotic"  bacteria  probably  serve  as  food  for  the 
amoebse,  which  need  solid  pabulum. 


34 


SPIROCILETES. 


spirochaetes  longitudinal  division  is  described  in  Sp. 
duttoni  by  Mayer  and  by  Newstead;  transverse  by 
Duval  and  Todd,  and  by  Koch.  In  Sp.  obermeieri,  (Fig. 
29),  both  longitudinal  and  transverse  division  are  found 
(Mackinnon;  Nuttall,  Fantham,  and  Porter).  In  Sp. 
pallida,  longitudinal  division  is  upheld  by  Beer  and  by 


FIG.  28. — Spirochaeta  anodontae;  transverse  division.     (Schellack.) 

Krzystalowicz  and  Siedlecki  (Figs.  30,  31);  transverse 
by  Levaditi  and  Yamamouchi.  In  Sp.  vesperuginis, 
Nicolle  and  Compte  observed  transverse  division, 
Gonder  on  the  other  hand  longitudinal  (Fig.  32).  In 
Sp.  gallinarum,  Borrel  saw  transverse  division,  Prowa- 
zek,  longitudinal  (Fig.  33) ;  the  latter  method  is  described 


GENERAL   CHARACTERS   OF   SPIROCELETES. 


35 


by  Gonder  in  Sp.  hartmanni,  the  former  by  Martin  in 
the  spirochaete  of  the  horse  and  by  Wenyon  in  that  of 
the  mouse.  Forked  forms  which  are  not  infrequently 


Ji 

FIG.  29.  FIG.  30.  FIG.  31.  FIG.  32. 

FIG.  29. — Spirochaeta  recurrentis  (Sp.  obermeieri);  transverse 
division.  (Schellack.) 

FIG.  30. — Longitudinal  division  of  Spirochaeta  pallida.  (Krzys- 
talowicz  and  Siedlecki.) 

FIG.  31. — Spirochaeta  pallida;  multiple  transverse  fission. 
(Krzystalowicz  and  Siedlecki.) 

FIG.  3  2 . — Spirochaeta  vesperuginis;  transverse  division.  (Gonder.) 


FIG.  33.  FIG.  34. 

FIG.  33. — Spirochaeta  gallinarum,  longitudinal  division:  a, 
First  stage;  b,  second  stage  (transverse  fission  ?) ;  c,  daughter  form 
after  division.  (Prowazek.) 

FIG.  34. — Spirochaeta  obermeieri;  filamentary  forms  in  the  bug. 
(Klodnitzky.) 

observed  are  held  to  point  to  longitudinal  division, 
forms  with  attenuated  central  strand,  to  transverse 
multiplication ;  but  the  latter  formation  has  been  held  by 


3  6  SPIROCILETES. 

the  supporters  of  longitudinal  division  to  represent 
the  final  stage  in  this  process,  just  before  separation  of 
the  daughter  individuals  takes  place  (Fig.  33). 

Gross  believes  that  spirochaetes  become  bent  upon 
themselves  before  undergoing  transverse  division,  the 
close  apposition  of  the  two  limbs  of  the  bent  organism 


FIG.  35.  FIG.  36.  FIG.  37. 

FIG.  35. — Spirochasta  balbianii ;  supposed  conjugation.     (Perrin.) 
FIG.  36. — Spirochaeta    vesperuginis;    conjugation   (?).     X22$o. 

(Gonder.) 
FIG.  37. — Spirochaeta  anodontae;  apparent  conjugation,  probably 

a  stage  in  longitudinal  fission. 

then  giving  rise  to  an  appearance  of  longitudinal  fission. 
He  terms  the  process  division  by  "incurvation." 

A  development  of  the  spirochaetes  into  long  fila- 
ments has  been  described  by  Duval  and  Todd  in  culti- 
vations of  Sp.  duttoni,  and  by  Klodnitsky  as  occurring 
in  Sp.  obermeieri  within  the  body  of  the  bug  (Fig.  34). 
Koch  also  saw  filamentary  forms  in  ticks  infected  with 
Sp.  duttoni. 


GENERAL   CHARACTERS    OF    SPIROCH^TES.  37 

The  occurrence  of  conjugation  in  some  form  or  other 
appears  to  be  universal  among  protozoa,  and  if  spiro- 
chaetes  are  to  be  assigned  to  this  class  it  will  presumably 
be  proved  to  take  place  among  them  also.  At  present 
the  accounts  given  of  such  an  occurrence  are  not  con- 
clusive (Figs.  35,  36).  An  instance  which  I  observed 
of  what  appeared  to  be  conjugation  in  Sp.  anodontcz 
(Fig.  37)  was  probably  only  a  stage  in  the  process  of 
longitudinal  division.  Krzystalowicz  and  Siedlecki 
have  explained  certain  forked  forms  of  Sp.  pallida 


FIG.  38.  FIG.  39. 

FIG.  38. — Spirochaeta  pallida;  formation  of  gametes.  (Krzys- 
talowicz and  Siedlecki.) 

FIG.  30. — Spirochaeta  pallida;  conjungation.  (Krzystalowicz 
and  Siedlecki.) 

which  they  observed  as  resulting  from  the  conjugation 
of  a  small  individual  with  a  larger  one  (Figs.  38,  39) ; 
and  Leuriaux  and  v.  Geets  describe  microgametes  and 
macrogametes.  The  four  writers  last  mentioned  agree 
in  considering  Sp.  pallida  to  be  in  reality  a  species  of 
trypanosome  (Trypanosoma  luis,  Krz.  and  S.),  the 
macrogamete  being  definitely  trypanosomatoid  and  the 
microgamete  spirochaetal.  Male,  female  and  indifferent 
forms  of  Sp.  balbianii  are  described  by  Perrin,  but  he 
failed  to  find  any  certain  instance  of  conjugation  (see 

Fig-  35)- 

Carter  gives  figures  of  what  appears  to  be  conjuga- 
tion of  a  spirochaete  which  he  found  in  human  blood  in 


SPIROCILETES. 


S.  Arabia.  Large  globular  swellings  were  formed  in 
the  bodies  of  the  organisms  and  fusion  seemed  to  take 
place  between  two  individuals  at  these  points  (Fig.  40) . 


FIG.  41. 


FIG.  40. 

FIG.  40.  —  Arabian  spirochaste;  conjugation. 
FIG.  41.  —  Spirochaeta  balbianii;  encystment. 


(Carter.) 
(Perrin.) 


The  forms  depicted  are,  however,  so  unlike  those  seen 
in  any  other  spirochaetes  that  it  seems  possible  that 
Carter  was  dealing  with  a  spirochsetoid  form  of  some 
other  organism. 


FIG.  42. 


FIG.  43. 


FIG.  44. 


FIG.  45. 


FIG.  42. — Spirochaeta  anpdontae,  rolled  up  forms.     X  500. 

FIG.  43. — Spirochaeta  spiculifera,  rolled  up.     (Schellack.) 

FIG.  44. — Spirochaeta  pallida.  Depression  stage.  X2250. 
(Prowazek.) 

FIG.  45. — Resting  stage  of  Spirochaeta  buccalis.  X2250. 
(Prowazek.) 

Encystment  was  believed  by  Perrin  to  take  place  in 
Sp.  balbianii,  and  he  gives  illustrations  of  the  movements 
of  the  organism  in  assuming  this  phase  (Fig.  41).  The 


GENERAL   CHARACTERS    OF   SPIROCELETES.  39 

figures  which  he  gives  of  the  encysted  stage  are  not  very 
convincing.  In  the  case  of  Sp.  anodonta  I  have  found 
rolled  up  specimens  after  keeping  the  organisms  for 
a  day  or  two  in  the  fluid  which  escapes  from  the  mantle- 
cavity  of  the  mussel  (Fig.  42).  These  forms  in  some 


FIG.  46.— Spirochaeta  anodontae;  stages  in  formation  of  resting 
spore  (?). 

instances  lay  in  or  in  close  apposition  to  large  epithe- 
lioid  cells ;  but  whether  the  spirochaetes  entered  the  cells 
at  this  stage,  or  the  cell  engulfed  the  spirochaete,  or  the 
apposition  was  merely  due  to  some  physical  attraction, 
I  cannot  say  (compare  Figs.  69,  86,  87).  The  cells  were 


V 


FIG.  460. — Portion   of   Cladothrix   putridogenes    (Veszpremi)  for 
comparison  with  Fig.  46. 

necessarily  somewhat  degenerated  after  being  kept  so 
long  in  the  fluid,  and  it  seems  unlikely  that  they  would 
have  been  able  to  engulf  the  spirochaetes. 

Rolled  up  specimens  of  Sp.  duttoni  were  seen  by 
Mayer,  and  of  Sp.  pallida  by  Levaditi  and  others. 


40  SPIROCILETES. 

Oval  forms  were  found  by  Leuriaux  and  v.  Geets 
in  Sp.  pallida;  they  suggest  that  these  develop  subse- 
quently into  filaments.  Ovoid  bodies  are  also  depicted 
by  Krzystalowicz  and  Siedlecki  as  apparently  constitut- 
ing a  stage  in  Sp.  pallida  (see  Fig.  50),  and  somewhat 
similar  bodies  were  found  by  Castellani  along  with  Sp. 
pertenuis. 

Sporulation. — In  the  case  of  Sp.  anodontce  I  have 
found  some  specimens  showing  about  the  middle  of  the 
body  a  rounded  swelling  or  a  darkly  staining  oval  body 


FIG.  47.  FIG.  48. 

FIG.  47. — Spirochaeta  balbianii,  with  central  swelling  (so-called 
"  male"  form).     (Perrin.) 
FIG.  48. — Arabian  spirochaete.     (Carter.) 

(Fig.  46)  suggestive  of  a  "resting  spore."  In  figure 
460  is  depicted  for  comparison  a  portion  of  a  filament 
of  cladothrix  showing  a  very  similar  swelling.  These 
appearances  may  be  compared  with  the  illustrations 
given  by  Perrin  of  peculiar  forms  of  Sp.  balbianii  (Fig. 
47),  and  also  Carter's  figures  of  his  Arabian  spirochaete 
(Fig.  48). 

Cox  states  that  within  the  blood-corpuscles  Sp. 
obermeieri  breaks  up  into  minute  granules;  and  Leish- 
man  finds  that  Sp.  duttoni  gives  rise  to  coccoid  forms 
within  the  tick,  no  spirochaetes  being  discoverable  within 
ticks  which  are  yet  capable  of  conveying  the  infection. 
The  so-called  "moniliform"  specimens  of  spirochaetes 
(Fig.  27),  are  usually  looked  upon  as  degenerative  forms 
but  it  is  possible  that  the  granules  thus  formed  are  cap- 
able of  subsequent  development,  and  are  the  equivalent 


GENERAL   CHARACTERS    OF    SPIROCTLETES.  41 

of  bacterial  spores.     The  terminal  nodules  of  the  smaller 
spirochaetes  are  very  suggestive  of  such  a  formation. 

Associated  with  Sp.  anodonta  are  found  chains  of 
coccoid  bodies,  similar  in  length  to  the  spirochaetes. 
The  arrangement  of  the  staining  material  in  the  bodies 
of  the  latter,  which  may  break  up  into  regular  portions 


FIG.  49. — Development  (?)  of  Spirochaeta  anodontae:  a,  Spi- 
rochaete  apparently  undergoing  segmentation;  b,  chain  of  cocci, 
possibly  resulting  from  this  process  of  fission. 

of  about  equal  size,  suggests  that  they  may  give  rise  to 
the  cocci  by  a  process  of  multiple  fission  (Fig.  49) . * 

In  this  connexion  it  is  interesting  to  note  the  occur- 
rence of  rod-shaped  bodies,  containing  dots  resembling 
nuclei,  in  association  with  different  kinds  of  spirochaetes. 
Thus  they  were  found  by  Krzystalowic  and  Siedlecki 
along  with  Sp.  pallida  (Fig.  50)  by  Prowazek  in  connex- 

1 1  hope  to  publish  shortly  further  observations  supporting  the 
probability  of  this  mode  of  development  in  Sp.  anodontce. 


42  SPIROCH^ETES. 

ion  with  Sp.  gallinarum  (Fig.  51)  and  with  Sp.  lutra 
(see  Fig.  71),  by  Mayer  in  the  blood  of  a  mouse  infected 
with  Sp.  duttoni  (Fig.  52),  and  by  other  observers  else- 
where. Loewenthal  found  rod-shaped  and  sausage- 


FIG.  50.  FIG.  51. 

FIG.   50. — Abnormal  forms  associated  with  Spirochseta  pallida. 
(Krzystalowicz  and  Siedlecki.) 
FIG.   51. — Spirochaeta  gallinarum  and  rod  forms.     (Prowazek.) 

shaped    bodies    along  with    spirochaetes   in   ulcerated 
cancers. 

Qu£ry  believes  that  Sp.  pallida  is  a  stage  in  the 
development  of  a  bacillus  which  he  cultivated;  from 
it  he  prepared  a  serum  which  Hallopeau  found  to  have 


FIG.  52. — Rod  forms  (a,  c)  and  Spirochaeta  duttoni  (6)  in  blood 
of  mouse.     (Mayer.) 

some  curative  value  in  syphilis.  Von  Niessen  also 
holds  that  the  Sp.  pallida  is  a  developmental  form 
of  a  special  cocco-bacillus  which  he  obtained  in  pure 
culture. 

ASSOCIATION  WITH  OTHER  ORGANISMS. 

The  association  of  spirochaetes  with  other  organisms 
is  of  considerable  interest  as  either  constituting  an 
example  of  symbiosis  or  suggesting  the  possibility  that 


GENERAL   CHARACTERS    OF   SPIROCH^TES.  43 

these  organisms  are  a  stage  in  the  development  of  some 
other  form. 

The  best-known  example  of  such  constant  associa- 
tion is  that  of  the  spirochaste  met  with  in  Vincent's  an- 
gina with  the  fusiform  bacilli  which  are  often  regarded 
as  the  cause  of  this  condition.  The  idea  that  the 
two  forms  may  be  stages  in  the  life-history  of  one  and 
the  same  organism  is  attractive  and  is  supported  by 
the  experiments  of  Ruth  Tunnicliffe  who  found  that 
in  cultures  of  B.  fusiformis  spirochsetes  made  their 


FIG.    53. — Spirochaeta  vincenti  (a)  with  fusiform   bacilli  (6)  and 
(?)  filament  of  cladothrix  (c). 


appearance.  This  observation  has  not,  however,  been 
confirmed.  Schmiedlechner  found  bacilli  which  di- 
vided longitudinally  along  with  Vincent's  spirochaetes. 
Organisms  resembling  B.  fusiformis  were  found  by 
Mayer  along  with  Sp.  duttoni  and  by  Krzystalowicz 
and  Siedlecki  with  Sp.  pallida;  and  Leishman,  Harvey 
and  Bousfield,  and  Launois  and  Loederich  noted  the 
coexistence  of  Sp.  pallida  with  fusiform  bacilli  in  syphi- 
litic lesions.  Wellmann  found  Sp.  pertenuis  along 
with  fusiform  bacilli  and  with  Sp.  refringens. 

Other  bacterial  forms  are  also  found  associated  with 
spirochaetes,  as  by  Button,  Todd  and  Tobey  in  onychia, 
by  Schereschewsky  in  syphilis  (B.  pyocyaneus),  by 


44  SPIROCILETES. 

Cleland  in  castration-tumours  in  pigs,  and  by  Cleland 
and  by  Bosanquet  in  ulcerative  granuloma.  Levaditi 
found  it  impossible  to  cultivate  Sp.  pallida  and  Sp. 
rejringens  in  collodion  sacs  in  the  peritoneal  cavities 
of  animals  without  the  simultaneous  development  of 
bacterial  forms. 

In  gangrenous  stomatitis  the  spirochaetes  are  found 
most  deeply  situated  in  the  tissues,  the  bacilli  in  the 
more  superficial  parts;  while  in  ulcerative  granuloma 
bacilli  and  spirochaetes  are  found  together  deeply  situ- 
ated in  the  tissues,  where  it  would  seem  unlikely  that 
merely  saprophytic  germs  would  be  able  to  establish 
themselves.  The  bacteria  do  not  here  resemble  any 
known  pathogenic  variety. 

The  coexistence  of  trypanosomes  with  spirochaetes 
was  noted  by  Theiler  in  cattle,  by  Wenyon  in  mice,  and 
by  Petrie  in  birds  (martins).  Balfour  found  spiro- 
chaetes in  intestinal  ulcers,  in  dogs  and  monkeys,  pro- 
duced by  inoculation  with  trypanosomes  (7".  dimorphon); 
but  as  spirochaetes  or  spirilla  are  common  in  the  ali- 
mentary canals  of  animals,  they  may  have  been  only 
accidentally  present  in  these  cases.1  Gauducheau 
noted  the  appearance  of  spirillar  bodies  in  a  culture  of 
amoebae.2 

The  question  as  to  the  relationship  between  spiro- 
chaetes and  other  organisms  must  at  present  be  left 
undecided.  In  view  of  observations  recorded  in  this 
and  in  the  preceding  section  the  possibility  that  spiro- 
chaetes are  only  one  stage  in  the  development  of  a  poly- 
morphic organism  must  not  be  lost  sight  of  as  a  probable 
explanation  of  this  association,  though  it  is  at  present 
far  from  proof. 

Balfour,  Jour.  Trap.  Med.,  1906.  2 Gauducheau,  C.  R.  Soc.  de 
BioL,  1908,  LXIV,  493;  Leishman,  Jour.  R.  Army  Med.  Corps 
1905,  IV,  321;  Petrie,  Jour,  of  Hygiene,  1905,  V,  195. 


GENERAL   CHARACTERS   OF   SPIROCILETES.  45 

PATHOGENICITY. 

Since  it  has  been  found  impossible  to  produce  pure 
cultures  of  any  spirochsetes,  rigorous  proof  of  their 
causal  connection  with  disease,  such  as  is  demanded 
by  the  well-known  "postulates"  of  Koch,  cannot  be 
furnished.  Nevertheless,  the  pathogenicity  of  certain 
forms  appears  to  be  fairly  established. 

RELAPSING  FEVER. 

In  the  case  of  relapsing  fever  the  spirochaete  dis- 
covered by  Obermeier  and  generally  known  by  his  name 
is  universally  admitted  to  be  the  exciting  cause  of  the 
disease.  It  is  present  in  all  cases  of  the  malady;  it 
disappears  during  the  remissions  and  reappears  during 
the  relapses.  The  blood  of  a  patient  reproduces  a  similar 
affection  when  injected  into  animals,  and  no  other  germ 
has  been  found  present  which  might  be  looked  upon  as 
pathogenic. 

The  disease  is  characterised  by  a  sudden  onset,  after 
an  incubation-period  of  varying  length  (from  a  few 
hours  up  to  fourteen  days).  The  first  access  of  fever 
lasts  about  a  week  and  ends  by  crisis.  A  period  of 
apyrexia  ensues,  to  be  followed  by  a  second  access  of 
fever,  and  this  sequence  may  be  repeated  a  third  and 
even  more  times.  The  spleen  and  liver  are  much  en- 
larged, and  the  spirochaete  may  be  found  in  the  spleen 
during  the  remissions.  Monkeys  are  the  only  lower 
animals  directly  susceptible  to  the  disease. 

In  mankind  infection  may  be  conveyed  by  direct 
inoculation,  as  by  a  scratch  at  a  necropsy;  it  may 
also  occur  by  aerial  conveyance  in  some  instances,  like 
the  infection  in  scarlet  fever,  typhus,  etc.1  It  has 
been  supposed  that  the  infection  may  be  conveyed 

1  Ker,  "Infectious  Diseases"  London,  1908,  p  229. 


46  SPIROCILETES. 

from  person  to  person  by  the  bites  of  bugs,  and  it  is 
probable  that  at  least  accidental  infection  may  thus 
he  produced,  as  Tictin  found  the  spirochaete  present 
in  these  insects  and  infected  monkeys  by  injection 
of  their  macerated  bodies;  but  it  is  not  proved  that 
an  intermediate  host  is  necessary  or  that  any  develop- 
ment of  the  spirochaete  takes  place  in  the  bug.  Man- 
teufel  and  also  Sergent  and  Foley  believe  that  infection 
is  conveyed  by  lice  (Pediculi  capitis  et  corporis). 

The  blood  of  a  convalescent  from  relapsing  fever 
contains  substances  capable  of  destroying  the  spiro- 
chaetes  (spirillicidins) ,  and  passive  immunity  is  con- 
ferred by  injection  of  such  blood  into  another  indi- 
vidual. Agglutinating  substances  are  also  formed. 
Metchnikoff  holds  that  the  spirochaetes  are  mainly  de- 
stroyed by  phagocytosis;  Rabinowitsch  states  that  a 
hyperleucocytosis  is  produced  in  the  blood  of  animals, 
and  also  a  change  in  the  bone  marrow  "like  leukaemia." 

Relapsing  fever  has  been  chiefly  studied  and  described 
as  it  occurs  in  Europe  especially  in  Russia.  Closely  al- 
lied forms  of  disease  occur  in  other  parts  of  the  world, 
caused  apparently  by  spirochaetes  which  are  mor- 
phologically almost  indistinguishable  from  that  of 
ordinary  relapsing  fever,  but  are  probably  different 
species. 

AFRICAN  TICK  FEVER. 

African  tick  fever  is  an  affection  met  with  in  East 
Africa,  and  is  conveyed  by  the  bite  of  a  species  of  tick, 
Ornithodorus  moubata.  The  general  features  of  the 
disease  are  similar  to  those  of  relapsing  fever,  but  the 
relapses  are  often  more  numerous.  The  spirochaete 
present  in  this  disease  is  said  by  Uhlenhuth  and  Haendel 
to  be  rather  thicker  than  Sp.  obermeieri,  and  by  Schel- 
lack  to  move  more  vigorously.  An  attack  confers 


GENERAL   CHARACTERS    OF   SPIROCH^TES. 


47 


immunity  against  this  parasite,  but  not  against  Sp. 
obermeieri;  and  agglutination  tests  also  point  to  a 
specific  difference  between  the  two  spirochaetes.  Mice 
and  guinea-pigs  can  be  inoculated  with  this  organism, 
as  can  also  rabbits,  horses  and  monkeys.  The  patho- 
logical changes  seen  after  death  are  the  same  as  in 
infection  with  Sp.  obermeieri.  The  spirochaetes  are  taken 
up  by  the  ticks  from  the  blood  of  an  infected  person 
and  are  passed  on  in  the  egg  to  the  young  ticks  of  a 
second  generation.  Koch  describes  filamentary  forms 
of  the  spirochaete  in  these  animals.  Levaditi  and 
Manou^lian  believe  that  destruction  of  the  spirochaete 
and  cure  of  the  disease  are  brought  about  by  phagocy- 
tosis; some  of  the  symptoms  of  the  malady  are  attrib- 
uted by  them  to  thrombosis  of  blood-vessels  caused 
by  impaction  of  masses  of  spirochaetes. 


OTHER  FORMS  OF  RECURRENT  FEVERS. 

Relapsing  fever  as  seen  in  America  is  said  to  be  caused 
by  yet  another  spirochaete.  Schellack  gives  the  follow- 
ing description  of  the  three  allied  organisms: 


Length 

Breadth 

African  form  
(Sp.  duttoni) 
American  form  

24-3  OfJL 

(long  forms) 

o-45j« 

(Sp.  novyi) 
Russian  form  
(Sp.  obermeieri) 

(long  forms) 

19-20/1 

(long  forms) 

°-39^ 

It  is  not  impossible  that  other  species  may  be 
found  to  exist  in  different  parts  of  the  world  where 
forms  of  relapsing  fever  are  encountered.  Thus  the 


48  SPIROCaETES. 

spirochaete  of  relapsing  fever  as  seen  in  Bombay  was 
found  by  Novy  to  differ  from  the  American  variety. 

Spirochaetes  were  also  found  by  Carter  in  cases  of  fever 
occurring  of  S.  Arabia.  The  illness  consisted  in  head- 
ache, backache,  prostration  and  enlargement  of  the 
spleen.  Infection  appeared  to  be  conveyed  by  ticks 
(Ornithodorus  Sp.  ?).  The  relation  of  the  disease  to 
those  described  above  is  uncertain. 

Darling  believes  that  the  spirochaete  of  relapsing 
fever  in  Panama  is  again  a  special  variety,  the  immun- 
ity produced  by  an  attack  not  being  protective  against 
the  spirochaetes  of  the 'other  forms.  The  blood  is  infec- 
tive in  the  intervals  between  the  paroxysms.  Apes  and 
mice  are  susceptible  to  this  parasite. 

SYPHILIS. 

The  spirochaete  discovered  by  Schaudinn  in  the 
lesions  of  syphilis  and  first  named  by  him  Spirochcsta 
pallida  is  now  generally  accepted  as  the  cause  of  this 
disease.  Since  it  is  not  possible  to  obtain  the  organism 
in  pure  culture  and  to  use  such  cultures  for  the  inocula- 
tion of  animals,  rigorous  proof  of  causal  connection  can- 
not be  furnished.  It  is  held,  however,  that  as  the 
spirochaete  is  constantly  present  in  syphilis  (100  per 
cent,  of  cases,  Hoffmann  and  Prowazek)  and  is  not  found 
in  any  other  conditions,  it  has  as  much  claim  to  be 
regarded  as  the  infective  agent  as  has  the  Bacillus  leproz 
in  leprosy.  In  view  of  its  presence  in  the  initial  chancre, 
its  subsequent  passage  to  the  neighbouring  lymphatic 
glands,  its  presence  in  the  blood  in  the  secondary  stage 
of  the  disease,  and  the  occurrence  only  of  degenera- 
tive forms  in  tertiary  lesions  (gummas)  which  are  not 
supposed  to  be  infective,  the  features  of  the  disease 
may  be  satisfactorily  explained  as  due  to  this  organism. 
The  discovery  of  immense  quantities  of  spirochastes 


GENERAL   CHARACTERS    OF    SPIROCILETES.  49 

in  still-born  foetuses,  the  offspring  of  syphilitic  mothers, 
is  also  urged  as  a  strong  argument  in  favour  of 
the  infective  character  of  the  organism.  The  serum 
of  syphilitic  subjects  is  said  to  agglutinate  these 
spirochaetes. 

While  we  may  admit  that  a  strong  prima-facie  case 
can  be  made  out  for  this  view,  it  must  be  admitted  that 
there  are  still  some  objections  to  be  overcome  before 
the  Spiroch&ta  pallida  can  be  accepted  as  the  proved 
cause  of  syphilis. 

1.  In  the  absence  of  cultures  of  the  organism  it  can 
only  be   identified   by    its  morphological    characters. 
It  is  practically   admitted    that  only  those  observers 
who  have  made  a  prolonged  study  of  this  spirochaete 
and  of  similar  organisms  can  be  sure  of  distinguishing 
it  from  other  forms.     This  condition  of  affairs  some- 
what vitiates  the  statistical  method  of  proof,  dependent 
on  the  frequency  of  the  finding  of  Sp.  pallida  in  syphil- 
itic lesions. 

On  the  other  hand,  this  difficulty  of  identification 
may  be  used  as  an  argument  by  the  up-holders  of  the 
spirochaetal  hypothesis  against  those  who,  like  Kiolo- 
menoglou  and  Cube,  Krienitz  and  Scholtz,  have  found 
spirochaetes  which  they  identified  as  pallid®  in  non- 
syphilitic  conditions. 

2.  The  question  of  the  possible  variability  of  form  of 
Sp.  pallida  has  already  been  discussed   (page   30) ;  if 
such  variation  be   admitted,   it   becomes   practically 
impossible  to  be  sure  that  it  is  only  in  syphilitic  lesions 
that  this  spirochaete  is  found,  and  its  presence  elsewhere 
would  of  course  be  fatal  to  its  causal  connection  with 
syphilis. 

3 .  Most  observers  have  failed  to  find  the  spirochaete 
in  all  the  cases  of  syphilis  which  they  examined.     The 
following  figures  may  be  quoted : 


SPIROCH^ETES. 


Primary  lesion 

Secondary  stage 

Cases 

Posi- 
tive 

Nega- 
tive 

Cases 

Posi- 
tive 

Nega- 
tive 

Arning  and  Klein  

112 

108 

4 

179 

169 

10 

Bandi  and  Simonelli  

5 

3 

2 

Bodin  

16 

10 

6 

1  7 

10 

7 

Ferre"  

14 

1  1 

3 

J7 

17 

o 

Kraus  and  Prantschoff  .  .  . 

37 

32 

5 

25 

18 

7 

Metchnikoff 

6 

2 

Miihlens                 

22 

22 

o 

7 

6 

i 

Shennan  

10 

5 

5 

Siebert  

18 

13 

5 

47 

39 

8 

Bertarelli  and  Bovero  examined  forty-two  cases  and 
found  the  spirochaste  present  in  twenty-six,  absent 
in  sixteen.  They  noted  the  scanty  numbers  of  the 
organisms  which  could  be  found  in  the  primary  sore. 
Jaquet  and  Sevin  were  also  struck  by  the  difficulty 
of  finding  it  in  this  position.  Risso  and  Cipollina,  and 
also  Siebert,  record  the  inconstancy  of  the  organism 
in  syphilitic  lesions.  There  appears  to  be  agreement 
that  the  spirochsete  is  not  found  in  cases  of  "malig- 
nant syphilis"  (see  below).  This  is  curious,  since  the 
more  severe  the  disease,  the  greater  the  numbers  of  the 
pathogenic  organisms  which  should  theoretically  be 
present.  Even  if  these  cases  depend  rather  on  a  weak- 
ness of  resistance  on  the  part  of  the  patient  than  on  in- 
creased virulence  of  the  spirochaete,  there  should  still  be 
plenty  of  the  latter  present.1  The  possibility  that  the 
condition  is  due  to  a  mixed  infection  must,  however,  be 
borne  in  mind. 

The  relationship  of  the  spirochaetes  to  the  actual 
lesions  of  the  disease  presents  some  points  of  peculiar- 

^he  title  of  a  paper  by  Herxheimer  (Deut.  med.  Wochenschr., 
1905,  XXX,  1687)  suggests  that  he  found  Sp.  pallida  in  malig- 
nant syphilis,  but  the  paper  is  not  given  in  full. 


GENERAL  CHARACTERS   OF   SPIROCH^TES.  51 

ity.  It  would  be  natural  to  expect  that  in  so  well- 
marked  a  condition  as  the  primary  chancre,  which 
appears  to  be  produced  by  some  intense  irritant, 
there  would  be  found  large  numbers  of  the  causal 
organisms.  As  a  matter  of  fact,  spirochaetes  are  al- 
most invariably  scanty  in  the  tissues  forming  the 
chancre,  and  in  many  instances  observers  have  failed 
altogether  to  find  them.  (Their  presence  on  the  sur- 
face of  the  ulcer  is  of  comparatively  little  impor- 
tance, in  view  of  the  ubiquity  of  similar  organisms.) 
Others  have  failed  to  find  the  spirochaete  in  blood 
from  a  syphilitic  case,  which  was  definitely  proved  to 
be  infective.  The  possibility  of  the  existence  of  other 
stages  of  the  parasite,  e.  g.,  granules  or  coccoid  bodies 
(p.  40),  must,  however,  be  borne  in  mind. 

Again,  spirochaetes  may  be  present  in  the  organs 
of  a  syphilitic  patient  without  apparently  giving  rise 
to  any  anatomical  changes ;  as,  for  example,  in  an  eye 
which  appeared  quite  healthy  (Bab).  In  considering 
the  striking  abundance  of  spirochaetes  in  still-born 
syphilitic  foetuses  it  must  be  remembered  that  such 
bodies  may  be  found  full  of  other  (bacterial)  organisms, 
scattered  throughout  all  the  tissues;  and  also  that 
Karlinski  found  spirochaetes  distributed  throughout  the 
tissues  of  a  foetal  pig,  having  there  presumably  spread 
out  from  the  intestine  (see  Fig.  57). 

The  presence  of  the  spirochaetes  in  gummas,  which 
are  not  supposed  to  be  infectious,  is  somewhat  curious, 
for  while  this  non-infectivity  might  be  explained  on 
the  ground  that  the  spirochaetes  here  found  are  usually 
few  and  degenerate,  yet  if  they  are  the  cause  of  the  dis- 
ease at  this  stage,  they  must  be  of  very  notable  virulence, 
since  a  gumma  consists  of  a  considerable  mass  of  inflam- 
matory tissue  and  may  undergo  rapid  degeneration  and 
softening,  both  presumably  due  to  active  micro bial 


52  SPIROCILETES. 

action.     So  much  irritation  could  hardly  be  attributed 
to  the  presence  of  only  a  few  degenerate  spirochaetes. 

4.  The  fact  that  the  Sp.  pallida  is  found  in  the  lesions 
artificially  produced  by  inoculation  of  lower  animals 
with  syphilitic  material  constitutes  a  two-edged  argu- 
ment, since  the  only  animals  which  are  definitely  sus- 
ceptible to  syphilitic  infection  are  the  apes,  while  the 
spirochaete  has  been  found  living  in  dogs,  cats,  rabbits 
and  guinea-pigs.     It  is  necessary  to  assume  that  the 
pathogenic  agent  of  syphilis  can  exist  as  a  saprophyte  in 
lower  animals  which  are  insusceptible  to  the  disease — 
a  condition  which  is  at  least  unusual  in  the  case  of 
infective  organisms.     Otherwise  it  must  be  held  to  be 
saprophytic   in   both   cases.     In  this  connection  the 
observations  of  Gaylord  on  the  transmission  of  spiro- 
chastes in  mice  may  be  brought  into  comparison.     This 
observer  found  spirochaetes  in  certain  cases  of  cancer 
in  mice,  and  on  inoculating  other  mice  with  emulsions 
of  the  tumours,  he  found  that  the  spirochaetes  were 
also  discoverable  in  the  growths  which  resulted.     The 
natural  inference  that  the  spirochaetes  were  the  cause 
of  the  cancer  was,  however,  proved  to  be  erroneous; 
and  it  appeared  that  the  organisms  were  frequently 
present  in  mice  and  could  be  transmitted  from  one  to 
another,  but  were  harmless  parasites  of  these  animals. 
Of  course,  the  parallel  is  not  very  close,  since  the  Sp. 
pallida  is  not  found  in  normal  men  or  animals,  but  the 
possible  transmission  of  spirochaetes  which  are  in  no 
sense  pathogenic  must  be  noted. 

5.  The  discovery  of  spirochaetes,  very  like  Sp.  pallida 
in  the  closely  allied  diseases,  yaws  and  ulcerative  gran- 
uloma  of  the  pudenda,  would  seem  to  constitute  an  ad- 
ditional argument  in  favour  of  the  Sp.  pallida  being 
the  true  cause  of  syphilis.     Herein  also  lies,  however,  a 
certain  source  of  difficulty.     It  is  generally  said  that  the 


GENERAL   CHARACTERS    OF   SPIROCILETES.  53 

spirochsete  of  yaws  (Sp.  pertenuis)  is  indistinguishable 
from  Sp.  pallida.  So  far  as  the  silver- stained  prepara- 
tions made  by  Levaditi,  for  example,  are  concerned, 
this  is  true.  (On  the  other  hand,  Sp.  pertenuis  as  de- 
picted by  Castellani  is  different  in  appearance  from  Sp. 
pallida,  not  showing  the  same  regular  curls.)  If  then 
we  have  in  yaws  an  organism  exactly  resembling  Sp. 
pallida,  the  argument  that  this  organism  is  only  found 
in  syphilis  would  seem  to  break  down,  since  Sp.  pallida 
is  only  recognised  by  its  morphological  characters. 
The  identity  of  form  is  noted  by  Beurman  and 
Gougerot,  by  Levaditi  and  Nattan-Larrier,  by  Schiiffner 
and  by  Wellmann,  and  was  admitted  by  Schaudinn 
himself. 

6.  Levaditi  succeeded  in  cultivating  Sp.  pallida  (i.e., 
in  causing  some  multiplication  of  the  organisms  and 
keeping  them  alive)  in  collodion  sacs  placed  within  the 
peritoneal  cavities  of  monkeys.     These  cultures  failed 
to  produce  infection  when  inoculated  in  other  monkeys 
susceptible  to  syphilitic  infection.     The  negative  result 
is  not  perhaps  in  itself  a  very  strong  argument  against 
the  causality  of  the  spirochaete,  as  the  organism  may 
easily    have    become    attenuated    by  this   procedure. 
Nevertheless  it  may  be  pointed  out  that  Levaditi  allows 
that  a  similar  failure  in  the  case  of  the  spirochaete  of 
balanitis  is  evidence  against  the  causal  connection  of 
this  organism  with  this  disease,  and  the  argument  is  as 
strong  in  one  case  as  in  the  other.     A  positive  result 
from  inoculation  of  a  culture  has  since  been  recorded 
by  Bruckner  and  Galacesco. 

7.  The  argument  derived  from  the  destructive  and 
agglutinating   action   of   the  serum  of  syphilitic  pa- 
tients on  the  Spiroch&ta  pallida  is  somewhat  weakened 
by  the  fact  that  De  Lisle  and  Jullien1  found  a  coccus  in 

1  La  Semaine  Medicate,  1900,  page  279. 


54  SPIROCILETES. 

syphilitic  patients  which  was  agglutinated  by  their 
serum,  but  this  organism  is  not  now  supposed  to  have 
any  causal  connection  with  the  malady. 

8.  It  is  usually  said  that  the  result  of  antisyphilitic 
treatment  is  to  cause  the  disappearance  of  the  spiro- 
chsetes;  but  Rille  and  Vocquerodt,  and  also  Benda, 
found  that  no  apparent  effect  was  produced  by  treat- 
ment on  the  number  of  spirochaetes  present. 

9.  It  is  stated  that  whereas  the  syphilitic  virus  is 
active  when    infective    material   is  mixed   with  gly- 
cerine, spirochaetes  are  destroyed  by  contact  with  this 
reagent. 

After  a  review  of  all  the  evidence  at  present  available 
we  can  therefore  only  conclude  that  while  there  is  a 
considerable  probability  that  the  Spiroch&ta  pallida 
is  the  infective  agent  in  syphilis,  its  position  is  not  yet 
established  with  absolute  scientific  certainty. 

The  clinical  phenomena  of  syphilis  consist  in  (i)  a 
primary  lesion  at  the  site  of  inoculation,  consisting  in 
an  ulcer  with  a  deeply  indurated  base;  here  the  spiro- 
chaete  is  found  on  the  surface  of  the  ulcer  and  in  the  fluid 
that  can  be  squeezed  from  its  deeper  layers,  and  it  can 
also  be  found  by  the  silver-staining  method  lying  in  the 
lymphatic  spaces  between  the  bundles  of  fibrous  tissue. 
Enlargement  of  lymphatic  glands  takes  place  in  the  re- 
gion towards  which  the  lymph  from  the  ulcer  drains,  and 
by  puncture  of  these  glands  with  a  hypodermic  syringe 
the  organism  can  usually  be  obtained  in  the  fluid  with- 
drawn. (2 ) .  The  secondary  stage  is  marked  by  constitu- 
tional symptoms,  such  as  fever  and  feeling  of  illness, 
along  with  eruptions  on  the  skin  and  often  with  super- 
ficial ulceration  of  the  throat  (tonsils,  fauces).  In 
this  stage  the  spirochaetes  may  be  found  in  the  blood 
and  in  any  vesicular  lesions  which  may  develop,  or  in 
the  fluid  obtained  by  blistering.  (3).  In  a  certain  num- 


GENERAL  CHARACTERS   OF   SPIROCILETES.  55 

ber  of  cases  a  tertiary  stage  is  met  with,  consisting  in 
the  formation  of  localised  inflammatory  swellings, 
which  may  occur  in  any  part  of  the  body  (the  so-called 
gummas,  from  the  "gummy"  nature  of  the  material 
formed  by  breaking  down  of  the  inflammatory  cells). 
In  these  either  no  spirochsetes  are  found,  or  only  a  few 
degenerate  specimens.  This  stage  is  not  infective. 

Malignant  syphilis  is  the  name  applied  to  certain  forms 
of  the  disease  in  which  ulcerative  lesions  appear  at  an 
early  stage.  Anaemia  and  cachexia  develop  to  a  severe 
degree,  and  the  condition  is  not  amenable  to  treatment 
with  mercury.  It  "must  be  distinguished  from  severe 
syphilis  which  is  characterised  by  dangerous  localisa- 
tion of  the  disease,  or  by  association  with  other  morbid 
conditions.  The  cause  of  malignant  syphilis  is  doubt- 
ful; it  has  been  attributed  both  to  excessive  virulence 
of  the  microbe  and  to  secondary  infection  with  other 
organisms.  It  would  appear  more  probable  that  it  is 
due  to  the  implantation  of  the  microbe  on  virgin  soil — 
i.e.,  on  persons  whose  ancestors  have  been  free  from 
syphilis"  (C.  F.  Marshall1). 

Infection  is  usually  contracted  in  coitu  and  is  con- 
veyed by  the  discharge  from  the  primary  sore.  It  can 
also  be  conveyed  by  means  of  drinking  vessels  or  of 
a  tobacco  pipe  recently  used  by  an  infected  person 
in  the  secondary  stage,  in  which  there  are  infective 
lesions  in  the  mouth  and  throat.  Thus,  either  direct 
contact  is  necessary  or  very  direct  conveyance  by  an 
intermediary  object.  Some  abrasion  of  the  skin  is 
probably  required  for  the  entrance  of  the  infective 
organism,  and  surgeons  are  often  infected  through 
slight  wounds  in  the  fingers  in  the  course  of  examining 
syphilitic  patients.  Infection  does  not  seem  to  arise 
from  contamination  with  the  fluids  of  the  dead  body, 

1 "  Syphilology  and  Venereal  Disease,"  1906,  London,  pp.  18-19. 


56  SPIROCH^TES. 

as  in  making  post-mortem  examinations;  hence  it 
would  seem  that  the  organism  quickly  loses  its  viru- 
lence on  the  death  of  the  host. 

Congenital  Syphilis. — Infection  may  be  conveyed  to 
the  foetus  in  utero  from  either  parent.  Whether  the 
actual  spermatozoon  is  infected  cannot  be  demonstrated ; 
certainly  the  causal  organism  can  pass  from  the  mater- 
nal blood,  by  way  of  the  placenta,  into  the  vessels  of 
the  foetus.  In  this  way  congenital  syphilis  is  produced. 
The  syphilitic  foetus  often  dies  before  birth,  and  in  such 
still-born  foetuses  the  Spiroch&ta  pallida  is  found  in 
practically  all  the  organs  of  the  body — most  abundantly 
in  the  liver  (see  Fig.  88),  and  suprarenal  capsules.1 

It  is  generally  supposed  that  resistance  to  the  Sp. 
pallida  is  mainly  effected  by  phagocytosis;  but  Zabo- 
lotny  and  Maskalowetz  find  that  the  serum  of  syphilitic 
subjects  has  an  agglutinating  and  also  a  "lytic"  power, 
so  that  the  processes  at  work  in  producing  recovery 
and  immunity  may  be  complex  in  character.2 

Most  attempts  to  pass  the  virus  of  syphilis  through 
a  porcelain  filter  have  failed;  but  Jancke  records  one 

1  Some  of  the  actual  findings  are :    In  congenital  syphilis — in  the 
eye  and  nasal   mucus  (Bab),  in  the  blood  and  in  the  lungs  (De 
Sousa   and  Pereira),  in  the  bronchial  epithelium  and  in  miliary 
gummas  (Benda),  in  the  bullae  of  pemphigus   (Hoffmann,   Leixer, 
Levaditi),  in  the  testicles  (Fouquet),  in  osteitis  (Bertarelli),  in  the 
urine  (Huebschmann),  in  the  meconium  (Simmonds),  and  in  the 
placenta  of  the  mother  (Wallich  and  Levaditi).     In  the  acquired 
disease,  besides  its  localisation  in  the  connective  tissue  of  the  pri- 
mary sore,  and  in  the  lymphatic  glands  and  in  the  blood  in  the 
secondary  stage,  it  was  found  by  Ehrmann  in  the  nerve-sheaths, 
by  Follet  in  the  saliva.     Veillon  and  Girard  attribute  the  roseolar 
rash  to  the  formation  of   embolic  masses  of  the  spirochaetes  in 
the  blood-vessels.     The  organism  may  in  rare  instances  be  found 
in  the  cerebro-spinal  fluid  (Gaucher  and  Merle).     It  is  also  found 
in  aortitis  (Reuter,  Schmorl) .     The  spirochaetes  lie  as  a  rule  between 
the  cells  of  the  host;  but  they  may  be  seen  in  polymorphonuclear 
leucocytes   (Gierke)   where  they  may    be    undergoing    phagocy- 
tosis, and  also  inside  the  cells  of  the  liver  in  congenital  syphilis 
(Levaditi). 

2  The  formation  of  a  copula  or  amboceptor  has  already  been 
noted.     An  account  of  the  so-called  Wassermann  Reaction,  based 
on  this  occurrence,  does  not  fall  within  the  scope  of  this  little  work. 


GENERAL   CHARACTERS    OF    SPIROCILETES.  57 

positive   result   obtained   with   the   material   from   a 
syphilitic  foetus. 

YAWS. 

The  disease  called  yaws  or  pian  (Frambcesia  tropica) 
is  in  many  ways  like  syphilis,  going  through  a  primary 
stage  characterised  by  a  local  lesion  at  the  point  of 
inoculation,  and  a  secondary  stage  of  generalised 
phenomena,  which  take  the  form  of  multiple  nodular 
inflammatory  lesions  or  granulomas.  It  is  generally 
supposed  to  be  a  distinct  malady,  but  some  writers 
still  maintain  that  it  is  only  a  variety  of  syphilis.  The 
Sp.  pertenuis,  first  described  by  Castellani,  is  supposed 
to  be  the  causal  agent  of  the  disease.  It  so  closely  re- 
sembles Sp.  pallida  (page  112)  that  some  writers  are 
constrained  to  argue  from  the  diversity  of  the  two  affec- 
tions to  prove  the  distinct  characters  of  the  two  para- 
sites (e.g.,  Wellmann). 

Schiiffner  (who  believes  in  the  identity  of  yaws  and 
syphilis)  found  the  spirochaste  present  in  98  per  cent,  of 
all  the  cases  he  examined.  Von  dem  Borne  found  it  in 
seventy-three  of  seventy-six  non-ulcerated  cases:  in 
ulcers  it  is  found  along  with  other  kinds  of  spirochastes. 
Levaditi  and  Nattan-Larrier  found  the  organisms  on 
the  surface  of  the  primary  lesion,  in  the  glands,  and  in 
the  spleen,  but  not  in  the  blood.  Mclntosh  states  that 
the  position  in  which  this  spirochaete  is  found  in  the 
skin  is  not  the  same  as  that  in  which  Sp.  pallida  occurs ; 
Sp.  pertenuis  being  found  in  the  superficial  layers 
of  the  ulcer,  in  the  fibrinous  crust  containing  leucocytes 
and  degenerated  epithelium,  whereas  Sp.  pallida  occurs 
especially  in  the  corium  around  the  blood-vessels. 

LYMPHADENOMA. 

Proescher  and  White  discovered  spirochaetes  (Sp. 
lymphatica)  present  in  a  case  of  "  lymphosarcoma. " 


58  SPIROCILETES. 

With  material  containing  it  they  inoculated  a  monkey 
(Macacus  rhesus)  and  thus  produced  a  local  lesion  at  the 
point  of  inoculation,  followed  by  the  appearance  of 
other  nodules  at  a  distance.  They  further  inoculated 
a  second  monkey  from  the  first,  also  with  positive  re- 
sults. The  spirochaetes  were  found  in  all  the  lesions. 
These  spirochaetes  so  closely  resembled  Sp.  pallida 
that  they  were  at  first  mistaken  (?)  for  this  organism. 
It  seems  possible  that  in  this  case  either  a  syphilitic 
adenitis  was  mistaken  for  lymphadenoma,  or  that  a 
secondary  infection  with  Sp.  pallida  occurred  in  a 
patient  suffering  from  this  disease. 

ULCERATIVE  GRANULOMA. 

Spirochaetes  were  first  found  in  the  disease  called 
ulcerative  granuloma  of  the  pudenda  by  Wise,  who 
stated  that  some  forms  resembled  Sp.  pallida  and 
others  Sp.  pertenuis.  They  have  since  been  found  in 
this  disease  by  MacLennan  and  by  Cleland.  They 
exist  in  the  deep  layers  at  the  base  of  the  ulcer,  at  some 
distance  from  the  surface,  and  may  be  found  in  the 
blood-vessels.  Bacteria  are  present  along  with  the 
spirochaetes.  The  pathogenicity  of  these  organisms 
cannot  be  held  to  be  proved  as  yet,  but  their  position 
deep  in  the  tissues  is  against  a  purely  saprophytic 
character.  * 

VINCENT'S  ANGINA. 

Spirochaetes  are  found  along  with  fusiform  bacilli  in 
the  disease  called  Vincent's  angina,  and  also  in  some 
other  conditions  in  which  there  is  necrosis  of  tissue  (e.g., 

1 M.  Carter  states  that  he  has  found  in  a  case  of  this  disease,  pro- 
tozoon  parasites  in  the  form  of  minute  rounded  bodies  within  the 
cells.  He  believes  that  they  are  related  to  Leishmania  donovani 
(Lancet,  1910,  II,  1128). 


GENERAL   CHARACTERS   OF   SPIROCH^ETES.  59 

in  noma  or  gangrene  of  the  cheek,  in  hospital  gangrene, 
etc.).  Uffenheimer  thinks  the  spirochaetes  are  the 
pathogenic  agents  in  Vincent's  angina,  as,  if  the  tonsils 
are  removed  in  cases  in  which  these  organisms  are 
present,  further  ulceration  occurs.  Ellermann  also 
points  out  that  the  spirochaetes  are  found  more  deeply 
situated  in  the  tissues  in  cases  of  noma  than  are  the 
bacteria,  and  that  they  seem  to  prepare  the  way  for 
the  latter.  The  relationship  between  the  spirochaetes 
and  the  fusiform  bacilli  has  already  been  discussed 
(page  43).  Veszpr&ni  thinks  that  the  spirochaete  of 
Vincent's  angina  is  identical  with  the  common  spiro- 
chaete of  the  mouth,  Sp.  buccalis. 

OTHER    DISEASES    IN    MAN. 

Moritz  found  spirochaetes  in  the  bone-narrow  and  in 
the  muscular  coat  of  the  intestine  in  a  man  who  died 
with  symptoms  of  intense  anaemia,  fever  and  diarrhoea, 
and  in  whom  there  were  found  after  death  an  ulcer  of 
the  stomach  and  miliary  nodules  of  new  growth  (en- 
dothelioma?)  in  the  lungs.  The  primary  seat  of  the 
tumour  was  not  discovered.  The  spleen  was  enlarged. 
He  thinks  the  spirochaetes  in  this  case  were  different 
from  any  previously  described  species,  but  their  nature 
and  relation  to  the  disease  must  remain  problematical. 

Castellani  found  spirochaetes  present  in  certain  cases 
of  haemorrhagic  bronchitis  (Sp.  bronchialis) .  He  believes 
that  they  may  be  the  cause  of  the  disease. 

Waters  also  noted  spirochaetes  in  a  series  of  cases  of 
bronchitis,  without  haemoptysis.  Branch  found  or- 
ganisms of  this  nature  in  a  case  of  pulmonary  tuber- 
culosis, in  which  they  were  present  in  the  expectora- 
tion along  with  tubercle  bacilli;  they  may  have  been 
derived  from  the  mouth. 


60  SPIROCHjETES. 

Spirochaetes  are  also  supposed  to  be  responsible  for 
the  diseases  known  as  balanitis  and  ulcus  tropicum 
(see  pp.  107,  103). 

DISEASES  OF  ANIMALS. 

A  number  of  diseases  in  the  lower  animals  are  associ- 
ated with  the  presence  of  Spirochaetes.  The  best  known 
is  the  so-called  spirillosis  of  fowls,  caused  by  the  Sp. 
gallinarum.  The  disease  is  characterised  by  weakness, 
drowsiness  and  diarrhoea.  It  may  prove  fatal  or 
recovery  may  ensue.  The  anatomical  lesions  met  with 
consist  in  a  fatty  degeneration  of  the  liver,  with  accu- 
mulation of  uninucleated  cells  round  the  vessels,  and 
hyaline  necrosis  in  the  spleen.  The  spirochastes  are 
found  in  and  around  the  vessels,  in  their  walls,  and 
between  the  cells  of  the  liver.  They  do  not  usually 
become  intracellular,  but  may  be  found  in  the  mature 
ovules.  Hereditary  infection  does  not  occur,  the 
offspring  of  infected  fowls  being  immune.  The 
fertilised  egg  can,  however,  be  artificially  inoculated 
and  then  the  chick  is  born  infected.  In  such  chicks 
the  liver  is  first  invaded  and  undergoes  the  change  above 
described.  Haemorrhages  also  occur,  and  the  blood  is 
described  as  "myeloid"  or  "embryonic"  or  both  to- 
gether (containing  granular  myelocytes,  vacuolated 
uninucleated  cells,  and  nucleated  red  corpuscles  with 
basophile  granules) . 

The  Spirochaetes  are  agglutinated  by  the  serum  of  the 
infected  birds.  Levaditi  considers  that  they  are  de- 
stroyed by  phagocytosis,  carried  out  especially  by  the 
macrophages  of  the  liver  and  spleen.  Neufeld  and 
Prowazek  deny  this,  and  attribute  recovery  to  the 
action  of  the  serum,  which  is  parasiticidal  in  vitro  and 
also  capable  of  protecting  other  birds.  Gabritchewsky 
states  that  "lytic"  bodies  (bacteriolysins)  appear  in 


GENERAL   CHARACTERS    OF   SPIROCILETES.  6 1 

the  blood  before  phagocytosis  occurs  (in  the  goose), 
while  Levaditi  and  Lange  find  exactly  the  reverse,  the 
spirochastes  disappearing  before  the  formation  of  anti- 
bodies ("immobilisines")  in  the  rabbit.  Infection  is 
conveyed  by  the  bites  of  ticks  (Argas  miniatus,  A. 
reflexus.  A.  persicus,  Ornithodorus  moubata,  not  by 
Dermanyssus  avium).  Atoxyl,  a  drug  containing  ar- 
senic which  is  found  useful  in  the  treatment  of  trypano- 
somiasis,  is  also  valuable  in  fowl  spirochaetosis,  being 
both  preventive  and  curative.  Salvarsan  ("606")  is 
also  very  efficacious. 

A  disease  of  cattle  characterised  by  fever  and  diarrhoea 
with  enlargement  of  the  spleen  was  described  by  Thei- 
ler.  Crises  and  subsequent  relapses  may  occur.  Infec- 
tion is  conveyed  by  the  bite  of  the  tick  Rhipicephalus 
decoloratus,  and  it  would  seem  that  some  develop- 
mental stage  may  occur  in  this  host,  as  direct  injec- 
tion of  the  blood  of  an  infected  animal  into  a  healthy 
one  does  not  result  in  infection. 

Spirochaetes  were  also  found  by  Theiler  in  sheep  and 
horses  suffering  from  fever;  he  believes  that  these  may 
have  been  the  same  organism  as  was  found  in  the  cattle. 
Baruchello  and  Pricolo  found  Spirochaetes  in  cases  of 
infective  pleuropneumonia  in  horses,  the  organisms 
being  present  in  the  spleen,  in  the  pulmonary  alveoli, 
in  the  pleural  effusion,  and  in  the  blood.  Martin  also 
found  Spirochaetes  in  a  sick  horse,  which,  however,  re- 
covered, and  Stordy  in  a  horse  which  died  with  symp- 
toms of  wasting  and  oedema;  with  these  organisms  the 
latter  failed  to  infect  a  dog. 

A  disease  due  to  a  spirochaete  (Sp.  vesperuginis)  was 
described  in  the  bat,  Vesperugo  kuhlii,  by  Nicolle  and 
Compte.  Enlargement  of  the  spleen  was  found  in  a 


62  SPIROCH^ETES. 

fatal  case.     Recovery  by  crisis  and  subsequent  relapse 
were  noted. 

Dodd  found  spirochaetes  in  certain  lesions  of  the  skin 
in  a  pig,  infection  being  conveyed  by  contact.  The  or- 
ganisms disappeared  and  reappeared  during  the  course  of 
the  disease ;  they  did  not  get  into  the  blood  of  the  animal, 
and  inoculation  was  only  possible  in  the  skin.  The 
lesions  in  some  cases  healed,  yet  the  pig  died.  Anaemia 
and  pneumonia  were  found  post  mortem. 

Breinl  and  Kinghorn  noted  enlargement  of  the  spleen 
in  mice  infected  with  Sp.  laverani.  This  organism  may 
be  the  same  as  Sp.  muris,  described  by  Wenyon,  but  the 
dimensions  given  by  these  writers  are  not  identical,  and 
Wenyon  states  that  Sp.  muris  is  a  harmless  parasite. 
He  thinks  it  may  be  identical  with  Spirillum  minor 
(minus)  found  by  Carter  in  the  rat. 

SUMMARY  OF  THE  PHENOMENA  OF  THE  PATHOGENICITY 
OF  SPIROCH^ETES. 

Summarising  the  facts  just  recorded,  we  find  that 
there  is  a  group  of  spirochaetes  associated  with  affections 
in  which  the  symptoms  are  fever,  a  tendency  to  relapses, 
and  enlargement  oj  ike  spleen.  In  this  group  we  may 
place  the  spirochaetes  of  the  various  forms  of  relapsing 
fever,  the  spirochaetes  of  fowl-spirillosis,  and  perhaps 
those  found  in  the  bat.  In  these  diseases  the  spiro- 
chaetes give  rise  to  a  septicaemia — a  generalised  infection 
in  which  the  organisms  multiply  in  the  blood-stream — 
without  the  formation  of  any  local  lesion.  The  causal 
connection  of  the  spirochaetes  with  these  affections  is 
clearly  established. 

On  the  other  hand,  we  have  in  syphilis  and  yaws 
diseases  in  which  there  is  a  well-marked  primary  local 


GENERAL   CHARACTERS    OF   SPIROCILETES.  63 

lesion,  followed  by  a  generalised  infection.  Relapses 
do  not  occur  as  a  notable  feature,  and  enlargement  of 
the  spleen  is  also  inconstant.  The  causality  of  the 
spirochsetes  present  is  not  yet  absolutely  proved. 

In  a  third  group,  in  which  we  may  place  Vincent's 
angina,  ulcerative  granuloma  of  the  pudenda,  and  per- 
haps the  disease  recorded  by  Dodd  in  the  pig,  there  is 
merely  a  local  affection  of  the  skin  or  of  mucous  mem- 
brane, conveyed  in  the  last  two  cases  by  direct  contact. 

The  three  classes  may  be  considered  to  constitute  a 
descending  scale  of  virulence  on  the  part  of  the  para- 
sites, a  local  lesion  being  caused  when  the  degree  of 
virulence  is  slight,  a  local  lesion  with  subsequent  gen- 
eralisation when  it  is  rather  greater,  and  an  immediate 
septicaemia  when  it  is  most  intense.  Cases  of  congenital 
syphilis  in  which  death  occurs,  but  no  anatomical 
lesions  are  discovered,  have  been  attributed  to  a  septi- 
csemic  action  of  the  Sp.  pallida. 

METHODS  OF  STAINING. 

The  larger  spirochaetes  may  be  stained  by  the  ordi- 
nary methods  used  for  protozoa — haematoxylin  (Dela- 
field's  or  Heidenhain's),  Giemsa's  stain,  etc.  They  are 
not  stained  by  carmine — thus  resembling  bacteria 
rather  than  protozoa. 

For  the  smaller  spirochaetes,  especially  for  Sp.  pal- 
lida, a  large  number  of  methods  have  been  recom- 
mended. 

Giemsa's  Stain. — This  is  made  as  follows:  Azur  II 
eosin  3  grm.,  azur  II  0.8  grm.,  pure  glycerine  250  grm., 
methyl  alcohol  250  grm.  One  drop  of  this  solution  is 
added  to  i  c.c.  of  distilled  water.  Stain  for  ten  to  fifteen 
minutes  (see  Plate,  Fig.  A) . 

Schereshewsky  recommends  diluting  13  to  15  drops  of 
this  solution  with  10  c.c.  of  0.5  per  cent,  solution  of 


64  SPIROCIOETES. 

glycerine.  The  mixture  is  wanned  and,  if  no  precipi- 
tate occurs,  is  poured  on  to  the  specimen  and  left  for 
two  or  three  minutes. 

Schmorl  uses  Giemsa's  stain  for  sections  of  tissue, 
following  it  by  a  watery  solution  of  alum.  He  admits 
that  this  method  is  inferior  to  Levaditi's  silver  method. 

Forest  stains  first  with  Ziehl's  carbol-fuchsine  in  the 
cold,  which  dyes  all  spirochaetes  except  Sp.  pallida,  and 
then  stains  with  Giemsa's  stain  for  twelve  to  sixteen 
hours  in  the  cold  and  for  one-half  hour  at  70°  C. 

Foix  and  Mallein  note  that  the  staining  by  Giemsa's 
solution  may  be  hastened  by  the  use  of  an  electric 
current. 

Other  Methods. — Davidsohn  recommends  the  use  of 
cresyl  violet  for  staining  Sp.  pallida;  Oppenheimer 
and  Sachs  use  carbolic  gentian  violet  (concentrated 
alcoholic  gentian- violet  solution,  10  c.c.,  5  per  cent, 
solution  of  phenol,  100  c.c.).  Proca  and  Vasilescu  use 
Gino  de  Rossi's  cilia-stain  (dissolve  50  grm.  pure  phenol 
and  40  gr.  tannin  in  100  c.c.  water  and  add  to  this  2.5 
grm.  basic  fuchsine  dissolved  in  100  c.c.  absolute  alcohol. 
Stain  in  this  for  ten  minutes,  wash  and  dry.  Then 
stain  with  a  mixture  containing  concentrated  alco- 
holic gentian  violet  10  c.c.,  phenol  5  grm.,  distilled 
water  100  c.c.). 

Reitmann  advises  that  films  should  be  first  treated 
with  5  per  cent,  solution  of  phosphoric  acid  in  water 
for  five  minutes  and  then  stained  with  carbol-fuchsine, 
warmed. 

Goldhorn  uses  a  complicated  preparation  of  poly- 
chrome methylene  blue  (methylene  blue,  lithium  car- 
bonate and  eosin),  and  McNeal  also  uses  methylene 
blue  and  eosin  (crude  methylene  blue  20  parts,  pure 
medicinal  methylene  blue  10  parts,  eosin  (yellowish) 
20  parts,  and  pure  methyl  alcohol  100  parts.  Stain  on 


GENERAL   CHARACTERS    OF   SPIROCILETES.  65 

cover-slip  for  forty-five  to  sixty  seconds ;  wash  in  dilute 
solution  of  sodium  carbonate — i  drop  of  i  per  cent, 
solution  in  10  c.c.  water). 

Herxheimer  and  Hubner  advise  the  use  of  bleu  de 
Nil  or  bleu  de  Capri.  Modifications  of  Jenner's  or 
Romanowsky's  stain  are  used  by  Simonelli  and  Bandi 
and  by  Von  dem  Borne.  The  former  writers  advise  the 
method  of  May  Grunwald,  viz.,  dissolve  i  grm.  of  eosin 
(Grubler)  in  i  litre  distilled  water,  and  i  gr.  of  methyl- 
ene  blue  (Meister,  Lucius  and  Briining)  in  a  similar 
quantity  of  water;  mix  these  solutions,  allow  to  stand 
for  two  to  seven  days,  and  filter.  Dissolve  the  sedi- 
ment in  pure  methyl  alcohol,  and  stain  preparations 
with  this  solution  for  four  to  ten  minutes. 

Impregnation  with  Silver. — Bertarelli  and  Volpino 
first  introduced  this  mode  of  staining  spirochaetes  in 
tissue,  but  the  modification  of  their  method  devised 
by  Levaditi  is  usually  preferred.  His  original  method 
is  as  follows: 

Fix  fragments  of  tissue,  about  i  mm.  thick,  in  10 
per  cent,  formol  solution;  wash  in  water,  and  harden 
in  96  per  cent,  alcohol ;  then  place  in  distilled  water  till 
they  sink.  Next  place  in  the  silver  solution  (silver 
nitrate  1.5  to  3  per  cent.)  at  a  temperature  of  33°  C. 
for  three  to  five  days.  Wash  in  distilled  water,  and 
place  in  the  reducing  fluid  (pyrogallic  acid  2  to  4  per 
cent.,  formol  5  c.c.,  distilled  water  100  c.c.)  for  twenty- 
four  to  forty-eight  hours.  Wash  in  distilled  water, 
dehydrate,  embed  and  cut  (see  Plate,  Fig.  B).  A 
subsequent  staining  of  the  sections  with  Giemsa's  stain 
or  with  toluydene  blue  may  be  carried  out. 

A  subsequent  modification  of  this  method  was  de- 
vised by  Levaditi  and  Manouelian,  by  which  the  first 
solution  is  made  of  i  per  cent,  silver-nitrate  solution 
with  the  addition  of  10  per  cent,  pyridine;  the  tissues 
5 


66  SPIROCILETES. 

are  then  washed  in  pyridine  (10  per  cent.) ;  and  reduc- 
tion is  effected  by  a  fluid  containing  pyrogallic  acid, 
4  per  cent.,  10  per  cent,  acetone,  and  15  per  cent, 
pyridine  in  distilled  water. 

Ravaut  and  Ponselle  use  an  albuminate  of  silver 
("largine")  in  2  per  cent,  solution,  followed  by  5  per 
cent,  pyrogallic  acid ;  the  largine  bath  may  be  repeated 
a  second  time  after  the  pyrogallic  solution,  the  tissue 
being  washed  with  distilled  water  between  the  two  baths. 

Yamamoto  uses  a  5  per  cent,  solution  of  silver  ni- 
trate, followed  by  a  solution  of  2  per  cent,  pyrogallic 
and  i  per  cent,  tannic  acid.  He  prefers  embedding  in 
celloidin,  and  uses  LoefBer's  methylene  blue  as  a  counter- 
stain. 

INDIAN    INK   METHOD. 

A  special  method  of  identifying  these  and  other 
organisms  on  a  field  rendered  opaque  by  means  of 
indian  ink  has  been  devised  by  Burri.  For  this  purpose 
ordinary  indian  ink  is  diluted  with  water  (i  part  in  6 
or  i  in  10).  The  solution  is  sterilised  and  allowed  to 
stand  for  two  weeks,  the  supernatant  fluid  being  then 
ready  for  use.  A  loopful  of  suspension  of  the  organ- 
isms is  mixed  with  a  drop  of  the  ink-solution,  spread  on 
a  slide  and  allowed  to  dry.  The  spirochaetes  are  then 
easily  seen  as  colourless  spirals  on  a  dark  background. 
Some  writers  prefer  a  stronger  solution  of  the  ink  than 
that  given  above,  e.g.,  i  in  2  of  water. 

EXAMINATION    OF     LIVING    SPIROCHAETES. 

Dark  Field  Examination. — The  identification  of 
living  spirochaetes  of  the  small  varieties  is  most  easily 
effected  by  examination  on  a  "dark  field"  under 
the  microscope,  by  means  of  one  of  the  forms  of  par- 


GENERAL   CHARACTERS    OF    SPIROCILETES.  67 

aboloidal  reflector.  The  scrapings  of  tissue  can  be 
examined  in  physiological  saline  solution  or  in  the  fluid 
exuding  from  the  sore,  or  a  drop  of  blood  can  be 
mounted  and  sealed  so  as  to  exclude  the  air.  The 
movements  of  the  spirochaetes  render  them  easily 
visible.  It  is  preferable,  however,  to  add  a  drop  of  dis- 
tilled water  to  "the  fluid  containing  the  organisms,  as 
this  causes  them  to  swell  somewhat  without  losing  their 
motility:  they  are  thus  more  easily  recognisable. 

Intra-vitam  Staining. — Mandelbaum  stained  living 
Sp.  pallidaz  in  a  hanging  drop  by  adding  a  loopful  of 
Lceffler's  methylene-blue  solution  along  with  a  loopful 
of  decinormal  soda-solution. 

Meirowsky  makes  a  paste  of  methyl- violet  and  salt- 
solution,  and  rubs  it  into  the  (previously  cleaned)  sur- 
face of  a  syphilitic  chancre.  In  the  serum  which  exudes 
there  are  found  stained  specimens  of  Sp.  pallida  and  Sp. 
refringens.  (Certain  more  deeply  staining  dots  in.  the 
substance  of  the  organisms  he  regards  as  nuclei.) 
Crystal  violet  is  as  efficacious  as  methyl  violet.  Sp. 
dentium  may  be  stained  by  spreading  a  drop  of  fluid 
containing  the  organisms  on  a  cover- slip,  and  placing 
this  face  downwards  on  a  drop  of  concentrated  watery 
solution  of  neutral  red  on  a  warm  slide. 

PSEUDO-SPIROCH^TES. 

A  vigorous  controversy  was  at  one  time  waged  as  to 
the  reality  of  the  spirochaetes  seen  in  sections  of  tissue 
stained  by  the  silver  method,  but  there  is  now  little 
doubt  as  to  the  genuine  nature  of  these  organisms.  It 
may  be  conceded  that  the  novice  might  be  deceived  by 
appearances  presented  by  nerve  fibres,  by  spirals  of 
fibrin  or  elastic  tissue,  or  by  the  edges  of  cells  which 
take  up  the  silver  and  present  a  crinkled  appearance 
(Fig.  54) ;  but  those  who  have  seen  actual  spirochaetes 


68 


SPIROCILETES. 


stained  by  this  method  could  hardly  confuse  them  with 
such  objects  or  mistake  the  latter  for  them. 

It  is  not  often  that  a  doubt  arises  as  to  the  reality  of 


FIG.  56. 


FIG.  54.  FIG.  55. 

FIG.   54.  —  Pseudospirochaetes;  edges  of  cells.     (Baling.) 
tes  in  a  smear  fro 
(From  a  specimen  by  Professor  E.  A. 


ng. 

FIG.   55.  —  Pseudospirochastes  in  a  smear  from  the  alimentary 
canal  of  Glossina  palpalis. 


Minchin.) 

FIG.   56.  —  Free  flagellum  of  Trypanosoma  lewisi. 

spirochaetes  when  other  stains  are  used,  but  Professor 
E.  A.  Minchin  has  in  his  possession  a  film  made  from  a 
"smear"  of  the  contents  of  the  alimentary  canal  of 


FIG.   57. — Pseudospirochaetes  or  spirochaetes  in  a  pig  foetus. 
(Saling.) 

Glossina  palpalis  (the  tse-tse  fly)  in  which  are  spiral 
objects  which  appear  to  be  artifacts,  but  which  have 


GENERAL   CHARACTERS    OF    SPIROCELETES.  69 

almost  exactly  the  appearance  of  true  spirochaetes  (Fig. 
55) .  The  reason  for  regarding  them  as  artifacts  is  that 
none  were  seen  in  other  specimens  of  the  same  material 
and  that  their  arrangement — often  in  crosses  or  radiat- 
ing lines — is  peculiar ;  but  their  resemblance  is  so  exact 
that,  if  this  view  be  correct,  doubt  may  be  thrown  on 
many  descriptions  of  spirochaetes  given  by  other 
writers. 

Objects  somewhat  resembling  spirochaetes  are  the 
male  gametes  of  some  protozoa  (coccidium,  plasmodium) , 
the  flagella  of  trypanosomes  which  may  occasionally  be 
found  free  (Fig.  56),  and  attenuated  forms  of  these  last 
organisms  and  of  Herpetomonas  or  Leishmania.  The 
appearances  found  by  Karlinski  in  a  macerated  pig- 
fcetus  and  figured  by  Baling  (Fig.  57)  may  be  either  real 
or  simulated  spirochaetes. 

CLASSIFICATION. 

It  has  already  been  pointed  out  that  the  position  of 
the  spirochaetes  in  nature — whether  they  are  to  be  re- 
garded as  protozoa  or  bacteria — is  doubtful.  Their 
relations  as  a  group  are  almost  equally  undecided. 
The  resemblance  of  the  smaller  to  the  larger  spirochaetes 
is  not  very  close.  Indeed,  while  each  of  these  groups 
is  fairly  well  marked,  the  relationship  between  the  two 
is  almost  less  striking  than  that  of  the  smaller  organisms 
to  a  group  of  bacterial  organisms — the  large  spirilla. 
The  Sp.  obermeieri  was  originally  classed  with  these,  and 
it  seems  not  impossible  that  this  view  is  more  correct 
than  the  recent  classification  of  these  little  organisms 
with  the  spirochaetes. 

Certain  peculiarities  observed  in  the  Sp.  pallida 
led  Vuillemin  and  afterward  Schaudinn  to  separate  it 
from  the  other  small  spirochaetes  and  to  put  it  in  a  dis- 
tinct genus,  called  by  the  former  Spironema  and  by  the 


70  SPIROCH,ETES. 

latter  (as  this  name  was  already  occupied)  Treponema. 
The  characteristics  of  this  genus  were:  The  round 
shape  of  the  body  in  section,  as  opposed  to  the  flattened 
shape  of  the  spirochaetes ;  the  fixity  of  the  curls  exhib- 
ited by  the  organism;  the  absence  of  an  undulating 
membrane;  and  the  presence  of  terminal  filaments. 
Sp.  pertenuis,  from  its  close  resemblance  to  Sp.  pallida, 
was  afterward  placed  in  the  same  genus. 

Examination  of  the  qualities  on  which  this  distinc- 
tion is  founded  does  not  tend  to  strengthen  a  belief  in 
its  validity.  The  round  shape  of  the  organism  in  section 
may  be  admitted,  but  the  ribbon-like  shape  of  the  other 
small  spirochaetes  is  not  very  clearly  established.  The 
existence  of  an  undulating  membrane  is  doubtful  even 
in  the  larger  spirochaetes,  and  still  more  problematical 
in  the  smaller  group.  Terminal  filaments  may  be  found 
in  other  species  besides  Sp.  pallida;  and  finally  the  even 
curls  presented  by  this  spirochaete  may  at  times  be  ab- 
sent, as  is  admitted  even  by  Schaudinn  himself.  Nor- 
ris,  Pappenheimer  and  Flournoy  point  out  that  the  Sp. 
obermeieri  exhibits  all  the  characteristics  assigned  to  a 
treponema. 

Blanchard  gives  the  following  classification : 
A.  SPIRO BACTERIA.     Cohn,  1875  (Spirillaceae,  Mi- 
gula,  1890).     Curved  bacteria;  rigid;  exhibiting  trans- 
verse division. 

1.  Spirosoma,  Migula,  1900.     Curved  rigid  rod;  no 
flagella.    (Sp.  nasale,  Wibel,  1887.    Sp.  linguale, 
id.,  1888.) 

2.  Vibrio,  Ehrenberg,  1878.    (Microspira,  Schrotter, 
1886.)    Rigid,  flagellated  rods,  joined  end  to  end, 
no  spores.     (V.  cholera,  etc.) 

3.  Spirobacillus,  Metchnikoff,   1889.     Large  spiral 
bacteria;  lateral  flagella;  spores. 


GENERAL   CHARACTERS    OF    SPIROCH.ETES.  71 

4.  Spirillum,  Ehr.,  1833.  Spiral,  cylindrical  or- 
ganisms ;  blunt  ends ;  no  undulating  membrane ; 
spores ;  terminal  flagella  (many  species) . 

B.  TRYPANOSOMID^:.  Doflein,  1901.  Flagel- 
lated; flexible;  undulating  membrane;  longitudinal 
division;  not  cultivable. 

1.  Spirochata,  Ehrenberg,  1833.   (Spirochaete,  Cohn, 
1 83  5 .)    Thin,  spiral ;  flattened ;  undulating  mem- 
brane;  no  flagella;  filiform  nucleus  and  chro- 
matin  granules  (type,  Sp.  plicatilis). 

2.  Treponema,     Schaudinn,     1905.       (Spironema, 
Vuillemin,  1905).     Characters  as  stated  above. 
(Trep.  pallidum.) 

3.  Trypanosoma,  Gruby,  1844. 

Liihe  divides  spirochaetes  into : 

1.  True  spirochsetes,  including  Sp.  plicatilis,  bal- 
bianii,     buccalis,     dentium,     vincenti,     vaccines, 
refringens,   pseudo-pallida,   and   the  spirochaete 
of  dysentery. 

2.  Blood-spirochaetes,    including    Sp.    gallinarum, 
anserina,  obermeieri,  duttoni,  theileri. 

3 .  Treponema :     Tr.  pallidum  and  pertenue. 

Gross  who  believes  that  spirochaetes  are  closely  allied 
to  the  bacteria,  coins  a  new  name  for  the  whole  genus, 
Cristispira,  derived  from  the  crest  or  ridge  which  he 
believes  to  run  along  one  side  of  the  body  or  around  it 
(this  formation  having  given  rise  to  the  statement  that 
spirochaetes  possess  an  undulating  membrane). 


SECTION  II. 

SYSTEMATIC  DESCRIPTION  OF  SPECIES. 

The  various  spirochaetes  will  be  considered  here  in  the 
following  order : 

LARGE  FORMS. 

Spirochaeta  plicatilis. 

Sp.  balbianii. 

Sp.  anodontae  and  other  spirochaetes  in  shell-fish. 

Sp.  polyspira. 

SMALL  FORMS. 

(Spirilla?) 
Sp.  obermeieri. 

Sp.  duttoni  and  other  species  in  relapsing  fever. 
Sp.  pitheci. 
Sp.  anserina. 
Sp.  gallinarum. 
Sp.  lagopodis.     Sp.  lovati. 
Sp.  vesperuginis. 
Sp.  lutrae. 

Sp.  theileri  and  allied  forms. 
Sp.  suis. 
Sp.  laverani,  Sp.  muris,  Sp.  minor,  Spirochaetes  in 

mouse-cancer. 
Sp.  gondii. 

Sp.  culicis,  Sp.  minei. 
Sp.  gadi,  Sp.  pelamydis,  Sp.  jonesii. 
Sp.  hartmanni. 
Sp.  bufonis. 

72 


SYSTEMATIC   DESCRIPTION    OF    SPECIES.  73 

Sp.  buccalis,  Sp.  dentium,  Sp.  vincenti,  Sp.  gracilis. 

Sp.  schaudinni. 

Spirochaetes  in  gangrenous  processes,  haemoptysis, 

etc. 

Sp.  microgyrata. 

Sp.  in  small-pox  and  vaccine  lymph. 
Sp.  refringens,  Sp.  balanitidis. 
Sp.  pseudo-pallida. 
Sp.  pallida. 
Sp.  pertenuis. 

Sp.  lymphatica  and  forms  found  in  anaemia. 
Sp.  aboriginalis. 
Sp.  interrogans. 
Various  doubtful  spirochaetes  and  spirilla. 

LARGE  SPIROCHAETES. 

SPIROCH^ETA  PLICATILIS. 

(Ehrenberg,  1833.) 

This,  the  first  discovered  species  of  spirochsete,  is 
found  in  stagnant  fresh  water,  especially  along  with 
algae  of  putrefaction.  It  attains  a  length  of  200^,  but 
its  average  length  is  80/1  and  its  breadth  0.5;*  (Doflein).1 
In  shape  it  shows  a  double  series  of  spiral  curves,  a 
larger  series  producing  a  generally  undulating  appear- 
ance, and  a  smaller  series  imposed  upon  these  (Figs,  i, 
58) .  Its  extremities  are  rounded,  according  to  Biitschli  : 
but  Laptschinsky  thinks  that  they  are  normally  sharp 
and  that  blunt  ends  indicate  that  the  organism  has 
broken  into  pieces.  It  is  flattened  in  form,  like  a  ribbon. 
It  possesses  a  periplastic  sheath  and  contained  ento- 
plasm.  Along  the  central  axis  is  a  refringent  central 
rod,  which  stains  darkly;  granules  of  staining  sub- 
stance (nuclei)  are  arranged  along  this  rod.  This  spiro- 
chaete,  according  to  Butschli,  has  no  undulating  mem- 

1  Illustrations  of  the  organism  make  it  appear  much  broader. 


74 


SPIROCILETES. 


brane,  Schaudinn  who  attributed  to  it  such  a  membrane 
probably  having  mistaken  for  it  the  loose  periplastic 
sheath.  It  exhibits  screw-like  movements,  but  also 
creeping  and  feeling  (kriechend,  tastend)  movements, 
while  the  contraction  of  its  sheath  may  lead  to  a  thicken- 
ing and  thinning  of  the  body  at  different  places.  Its 
mode  of  multiplication  is  not  known,  but  it  has  been 
seen  to  break  up  transversely. 


® 


IG.   58. — Spirochaeta  plicatilis.      (Schaudinn.) 

The  most  recent  description  of  this  organism  is  due 
to  Zuelzer,  according  to  whom  it  is  round  in  section, 
with  a  central  axial  thread  of  elastic  substance  and  a 
simple  spiral  twist.  A  round  refringent  body  is  placed 
at  the  central  point  of  each  wave,  and  fine  granules  of 
volutin  are  also  present.  Sometimes  a  terminal  gran- 
ule is  seen  at  one  extremity,  from  which  the  axial 
filament  starts.  The  length  may  reach  500/1  and  the 
breadth  £  /*.  The  ends  may  be  rounded  or  sharp. 
Transverse  division  occurs,  either  two  or  more  frag- 


SYSTEMATIC   DESCRIPTION   OF   SPECIES. 


75 


ments  being  thus  produced.  A  streaming  of  protoplasm 
may  be  seen.  There  is  no  undulating  membrane  or 
distinct  periplast.  The  axial  thread  is  important  for 
classification.  Sp.  plicatilis  may  be  found  in  the  sea 
(Naples)  as  well  as  in  fresh  water.  The  organism  is 
dissolved  by  trypsin. 

As  Sp.  plicatilis  is  the  "type  species  "  and  other  spiro- 
chaetes  must  justify  their  inclusion  in  the  genus  by  their 


FIG.  59. — Diagrammatic  representations  of  Spirochaeta  plicatilis; 

a,  according  to  Zuelzer,  straight  central  rod  and  dots  in  the  curls; 

b,  after  Schaudinn,  central  rod  follows  the  curves,  dots  superposed 
on  it  at  random;  c,  after  Doflein,  double  series  of  curls  with  cor- 
responding dots  and  indications    of    central    rod;  d,    imaginary 
figure  showing  central  rod  corresponding  with  dots  in  the  diagram 


resemblance  to  this,  it  is  unfortunate  that  so  great  a 
diversity  obtains  between  the  descriptions  given  by 
various  writers.  Thus  the  organism  is  flat  (Schaudinn, 
Doflein) ,  round  (Zuelzer) ;  it  has  a  double  series  of  curls, 
smaller  and  larger  (Doflein) ,  a  single  series  (Zuelzer) ; 
its  axial  filament  follows  the  curves  of  the  organism 
(Schaudinn) ;  it  runs  in  a  straight  line  (Zuelzer) .  These 
differences  are  illustrated  in  the  accompanying  dia- 


76  SPIROCILETES. 

gram  (Fig.  59).  It  is  noteworthy  that  the  nuclei  (?) 
are  distinct  from  the  axial  filament,  whereas  in  Sp. 
balbianii  the  central  staining  rod  is  regarded  as  nuclear. 

Very  remarkable  is  the  conflict  of  opinion  between 
such  observers  as  Butschli  and  Schaudinn  as  to  the 
possession  of  an  undulating  membrane  by  so  large  an 
organism  as  this.  Its  creeping  and  feeling  movements 
are  also  noteworthy,  as  being  unlike  those  seen  in  other 
spirochaetes.  The  movements  of  swelling  and  thinning 
may  be  compared  with  the  "euglenoid"  movement 
seen  by  Plaut  in  Sp.  vincenti. 

It  has  been  pointed  out  to  me  by  Mr.  T.  P.  Collings, 
to  whom  I  am  indebted  for  the  preparation  of  the 
illustrations  in  this  little  work,  and  whose  experience 
in  drawing  from  the  microscope  is  very  extensive,  that 
the  series  of  dots  seen  in  the  illustrations  of  Sp.  plicatilis 
is  very  suggestive  of  the  existence  of  a  spiral  rod  or 
filament  seen  in  optical  section  in  the  tissues  of  the  ' '  cork- 
screw". Fig.  59,  d,  shows  such  a  rod  corresponding 
with  the  position  of  the  dots  in  the  diagram  above  it. 
The  vast  experience,  however,  of  the  authors  who  have 
described  Sp.  plicatilis  makes  it  unlikely  that  they 
would  have  failed  to  recognise  such  a  structure  if  it  had 
been  present. 

SPIROCELETA    DAXENSIS. 

(Cantacuzfene,   1910.) 

Cantacuzene  found  in  the  water  of  the  hot  springs 
of  Dax  (52°  to  56°  C.)  a  spirochaete  closely  resembling 
Sp.  plicatilis.  It  was  from  30  to  loo/*  in  length  by 
^  to  2%n  in  breadth,  flattened  in  form,  and  exhibited  a 
double  series  of  curls,  smaller  waves  being  superposed 
on  the  larger  undulations.  It  possessed  a  longitudinal 
chromatin  filament  and  was  narrower  toward  the  ex- 
tremities, but  these  were  not  actually  pointed.  The 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  77 

organism  creeps  about  on  the  surface  of  the  algae  present 
in  the  water.  Cantacuzfene  distinguishes  three  kinds 
of  individuals :  i ,  slender  forms  (£/*  in  breadth) ,  the  axial 
filament  of  which  did  not  extend  quite  to  the  extrem- 
ities; 2,  shorter,  thicker  forms  (i  to  i%p  in  breadth), 
exhibiting  no  axial  filament  and  sometimes  occurring  in 
small  masses;  and  3.  a  few  specimens  showing  a  central 
swelling,  some  2^  in  thickness,  in  which  all  the  chro- 
matin  was  collected,  though  the  rest  of  the  organism 
stained  darker  than  did  the  other  forms.  Sometimes 
two  swellings  were  seen,  one  to  the  right  and  the  other 
to  the  left  of  the  central  point. 

SPIROCH^ETA    BALBIANII. 

(Trypanosoma  balbianii,  Certes,  1882.) 

This  large  spirochsete  was  first  described  by  Certes 
in  1882,  though  it  had  been  previously  seen  by  Moebius. 
It  is  found  in  the  crystalline  style  of  the  oyster  (Ostrea 
angulata,  0.  edulis).  It  attains  a  length  of  100  to  i2o/x 
and  a  breadth  of  5/x  (extremes  26  to  zoo/*  in  length; 
3  to  5/1  in  breadth,  Perrin).  Perrin  describes  an  in- 
different form,  a  female  form,  and  male  and  female 
gametes.  Taking  the  so-called  indifferent  form  as 
typical,  the  general  proportions  being  those  described 
above,  the  organism  consists  of  a  periplastic  sheath  with 
fluid  protoplasmic  contents.  The  former  is  of  some- 
what firm  consistency,  retaining  its  form  when  the 
contents  have  escaped.  It  is  furnished  with  an  un- 
dulating membrane,  running  spirally  round  the  body 
(Fig.  2).  This  has  a  dark  peripheral  fibre  (border  fibre, 
randfibrille)  or  thickening  of  the  edge  of  the  membrane. 
There  is  a  dark  nodule  at  each  end  of  the  periplast. 

The  nucleus  consists  of  a  spiral  band  (karyosome) 
formed  of  achromatic  substance  with  bars  of  chromatin 


78  SPIROCTLETES. 

at  regular  intervals;  sometimes  the  bars  may    exist 
separately  without  the  connecting  band  (Figs.  7,  60). 

Multiplication  takes  place  by  longitudinal  division, 
and  may  be  very  rapid  in  suitable  conditions,  giving 
rise  to  the  appearance  of  very  attenuated  individuals, 
scarcely  visible  under  the  microscope.  Divi- 
sion of  the  undulating  membrane  precedes 
that  of  the  body  of  the  organism.  When 
fission  is  nearly  complete,  the  daughter 
individuals  remain  attached  by  a  strand  of 
periplast  for  some  time,  appearing  as  a  single 
individual  of  abnormal  length  with  a  thin 
portion  in  the  middle.  In  the  process  of 
division  the  nucleus  first  condenses  to  form  a 
straight  rod,  lying  in  the  central  axis  of  the 
animal.  This  rod  breaks  up  into  short 
segments,  shaped  first  like  bacilli  and  then  as 
dumb-bells;  these  are  next  broken  up  into 
little  spherical  masses,  which  take  up  posi- 
tions in  two  rows  and  finally  in  groups  of 
four.  Perrin  believes  that  the  total  number 
of  these  is  sixty-four.  When  fission  has  oc- 
Spirochseta  curred  and  the  new  individuals  are  formed, 
balbianii.  the  nucleus  of  each  reforms  a  spiral  band. 

(Schellack.)       ^  ^   1  ,  e 

Encystment  takes  place  by  a  senes  of 
movements  by  which  the  two  ends  of  the  spirochaete 
first  travel  alternately  up  and  down  its  length,  and  then 
the  whole  organism  rolls  up  into  a  coil  (Fig.  41).  The 
periplast  then  bursts,  and  the  entoplasm  escapes,  tak- 
ing a  spherical  form,  but  not  secreting  any  covering 
membrane. 

The  female  forms  are  described  as  larger  and  stouter, 
with  a  thicker  periplast.  They  have  a  smaller  undulat- 
ing membrane,  and  encyst  less  readily.  The  nucleus 
goes  through  a  somewhat  different  course  of  changes  in 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  79 

encysting.  Masses  of  chromatin  are  formed,  connected 
by  thin  threads.  These  divide  into  fragments,  of 
which  the  greater  number  degenerate,  but  two  remain 
and  conjugate  (autogamy) ,  subsequently  giving  rise  to 
the  female  gametes. 

The  male  gamete  is  distinguished  by  the  appearance 
of  a  rounded  swelling  of  the  entoplasm  in  the  middle 
of  the  body,  followed  by  longitudinal  division  of  the 
organism,  starting  simultaneously  at  both  ends.  A 
reduction  of  chromosomes  takes  place  in  the  central 
swelling,  thirty-two  remaining.  Presumably  the  result- 
ing individuals  conjugate  with  female  forms,  but  Perrin 
notes  that  he  was  able  to  observe  only  a  few  doubtful 
examples  of  cojugation  (Fig.  35). 

Fantham  confirms  Perrin's  account  of  the  general 
form  of  the  nucleus.  Further  confirmation  is  needed 
of  the  different  forms  distinguished  by  Perrin,  and  of 
their  true  meaning,  but  we  may  compare  Gonder's 
account  of  Sp.  pinnce.  It  is  noteworthy  that  Perrin 
found  that  not  all  these  spirochaetes  had  undulating 
membranes,  and  also  that  he  observed  little  difference 
in  the  movements  of  the  forms  with  it  and  those  with- 
out it.  He  does  not  accept  the  view  of  Laveran  and 
Mesnil  that  the  organism  is  furnished  with  a  sheath, 
rather  than  with  an  undulating  membrane,  but  agrees 
with  them  that  it  is  a  spirochsete,  though  he  describes 
it  as  a  trypanosome. 

SPIROCH^ETA  ANODONT^E. 

(Keysselitz,  1906.) 

This  large  spirochsete  was  discovered  by  Keysselitz 
in  the  fresh- water  mussel  (Anodonta  cygnea) ;  it  also 
occurs  in  Anodonta  mutabitis.  It  exists  in  the  crys- 
talline style,  and  may  also  be  found  in  the  liver  and  in 


80  SPIROCELETES. 

the  gastric  epithelium.  It  attains  a  length  of  130^  and 
a  breath  of  3  to  4^.  It  is  said  by  Keysselitz,  to  have  a 
flattened  form  and  either  sharp  or  blunt  extremities 
(Figs.  61,  62,  620).  Those  with  sharp  ends  show  peri- 
plastic  appendages  (periplastfortstaze).  The  periplast 
is  fibrillary  in  appearance,  and  there  is  a  granule  (basal- 
korn)  at  each  end  and  an  undulating  membrane.  The 
nuclear  chromatin  takes  the  form  of  balls,  irregular 
masses  and  elongated  bands;  it  may  collect  in  a  single 
strand  toward  the  centre  of  the  organism.  Keysselitz 


FIG.  61. — Spi'-ochseta  spiculifera.     (Schellack.) 

did  not  observe  a  spiral  arrangement  like  that  described 
by  Perrin  in  Sp.  balbianii,  but  admits  that  such  may 
exist.  Division  is  by  longitudinal  fission. 

Schellack  notes  that  the  border  fibre  of  the  undulat- 
ing membrane  is  of  variable  thickness,  tapering  toward 
the  end,  unlike  that  of  trypanosomes,  and  believes  that 
it  is  not  a  true  membrane,  but  an  artifact  due  to  sepa- 
ration of  periplast.  I  am  inclined  to  agree  with  this 
view,  as  I  find  that  the  better  the  preparation  is  fixed 
the  fewer  organisms  appear  with  this  formation  (see 
page  20).  The  nucleus,  according  to  Schellack,  consists 
of  a  series  of  bands,  not  of  a  spiral,  and  the  whole 


SYSTEMATIC  DESCRIPTION   OF   SPECIES. 


81 


organism  is  built  up  of  a  series  of  chambers  arranged 
longitudinally.  Rolled  up  forms  occur  which  may 
possibly  represent  cysts.  He  also  believes  that  multi- 
plication is  by  transverse,  not  by  longitudinal  division 
(Fig.  28),  but  I  have  seen 
a  spirochaete  apparently  in 
process  of  longitudinal 
fission. 

The  blunt  and  pointed 
forms  described  by  Keys- 
selitz  are  held  by  Schel- 
lack  to  constitute  different 


FIG.  62. 


FIG.  62 a. 


FIG.  62. — Spirochasta  anodontae;  specimen  with  blunt  extremi- 
ties; different  sizes. 

FIG.  620. — Spirochaeta  anodontse  (Sp.  spiculifera)  forms  with 
pointed  ends;  different  sizes. 

species ;  to  the  latter  he  gives  the  name  Sp.  spiculifera 
(Fig.  61).  He  describes  a  long  form,  from  39  to  50. 5// 
long  and  o .  9  to  i .  2ju  broad  (average,  46/1  by  i/x),  and  a 
sharp-pointed  form,  28  to  36^  long  and  0.7  to  i .  i/t 
broad  (av.  32^  by  0.9^).  In  my  own  experience  four- 
teen specimens  with  blunt  ends  gave  an  average  length 
of  exactly  IOQ/J.  (extremes,  130  and  76jw),  while  of  fifteen 
sharp-pointed  specimens  the  average  length  was  55.6^ 


82  SPIROCILETES. 

(extremes,  96  and  34^).  The  short  variety  tends  to 
be  rather  thicker  than  the  long.  I  find,  however, 
spirochaetes  differing  greatly  in  length  and  even  more 
remarkably  in  thickness,  which  it  would  be  difficult 
to  sort  out  into  two  distinct  species.  I  am  inclined  to 
regard  the  specimens  with  pointed  ends  as  having  re- 
cently divided.  Schellack  admits  that  it  is  curious  that 
two  forms  should  be  found  to  coexist  in  each  of  three 


FIG.  63 . — Small  spirochaetes  from  the  alimentary  canal  of  Anodonta. 

different  molluscs  (Anodonta,  Tapes,  and  Ostrea), 
and  there  does  not  seem  any  good  reason  for  making 
separate  species  of  these  slightly  divergent  forms  at 
present.  On  the  other  hand,  without  stronger  grounds 
we  are  not  justified  in  describing  them  even  as  sexual 
forms  of  the  same  organism. 

I  have  found  in  the  alimentary  canal  of  Anodonta 
minute  spirochaetes,  10  to  i2//  in  length  (Fig.  63)  which 
may  be  identical  with  Sp.  hartmanni  of  Gonder  or  with 
Sp.  pusilla  of  Schellack  (see  below) . 

SPIROCH^TES  OF  OTHER  MOLLUSCS. 

Schellack  has  examined  a  large  number  of  different 
shell-fish  and  found  spirochaetes  in  many  of  them,  as 
shown  in  the  following  list.  He  believes  that  the 
average  measurement  of  individuals  taken  just  after 
division  is  a  good  criterion  of  species.  He  depicts 
many  spirochsetes  with  a  line  of  separation  in  the 
middle  of  their  length,  pointing  to  transverse  division. 
I  have  not  myself  been  able  to  observe  such  specimens 
in  the  case  of  Sp.  anodontoz,  and  even  in  the  longest  in- 


SYSTEMATIC   DESCRIPTION   OF   SPECIES. 


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84  SPIROCILETES. 

dividuals  which  I  have  seen  no  sign  of  division  was  dis- 
coverable. I  have,  on  the  other  hand,  noted  the  exis- 
tence of  thin  forms,  which  look  as  if  they  were  formed 
by  longitudinal  fission,  as  was  noted  by  Perrin  in  Sp. 
balbianii.  Hence  the  above  criterion  does  not  seem 
adequate  for  the  differentiation  of  species.  Fantham 
describes  Sp.  balbianii  as  occurring  in  Tapes — which 
points  to  the  probable  identity  of  Sp.  tapetos  and  Sp. 
balbianii,  and  throws  further  doubt  on  Schellack's 
classification.  The  following  table  embodies  the  lat- 
ter's  observations  (see  also  Fig.  64) : 

SPIROCH^ETA  PINN.E. 

(Gonder,  1908.) 

This  organism  occurs  in  Pinna  squamosa  and  P. 
nobilis.  It  varies  from  10  to  60/1  in  length  and  in 
breadth  from  |  to  3/x.1  One  end  is  blunter  than  the 
other,  and  here  is  situated  a  blepharoplast.  An  undu- 
lating membrane  is  present.  The  nucleus  may  take 
the  form  of  a  single  rod  or  of  irregular  masses,  which 
are  sometimes  arranged  in  groups  of  four.  The  rod- 
form  is  uncommon  and  is  seen  in  very  motile  forms, 
which  Gonder  regards  as  possibly  male  elements,  the 
irregular  masses  then  characterising  female  or  indiffer- 
ent forms.  A  concentration  of  all  the  chromatin  into 
one  rounded  mass  was  sometimes  observed.  Such 
individuals  may  constitute  involution  forms.  En- 
cysted forms  also  occur. 

SPIROCH^ETA  MACTR^E. 

(Prowazek,   1910.) 

Found  in  the  crystalline  style  of  the  mollusc,  Mactra 
sulcataria  (Deshayes),  in  Japan,  this  organism  is  from 
45  to  70^  in  length  by  0.8  to  i/i  in  breadth.  It  has  an 
undulating  membrane. 

1  The  dimensions  of  this  organism  given  by  Schellack  are  some- 
what different  (see  table  above). 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  85 


FIG.  64. — Spirochaetes  from  mussels.  (Schellack.)  a,  Sp. 
spiculifera;  b,  Sp.  cardii  papillosi;  c,  Sp.  saxicavae;  d,  Sp.  tapetos 
(X  2250);  e,  Sp.  spiculifera;  /,  Sp.  pusilla;  g,  Sp.  pusilla;  h,  Sp. 
anodontae;  i,  Sp.  limae;  k,  Sp.  ostreae;  /,  Sp.  modiolas.  X  1000  (ex- 
cept d) . 


86  SPIROCH^TES. 

SPIROCH^TA    PECTINIS. 

(Cristispira  pectinis,  Gross,  1910.) 
Gross  found  spirochaetes  free  in  the  alimentary  canal 
of  the  mollusc,  Pecten  jacob&us.  They  were  72/1  long, 
and  1.5/1  thick,  round  in  section,  with  blunt  ends. 
They  had  a  ridge  or  comb  (kamm,  crista)  running  along 
one  side,  but  no  terminal  filaments.  They  seemed  to 
consist  of  a  series  of  chambers  placed  one  behind  the 
other,  like  a  filament  of  some  alga. 

SPIROCH^ITA    INTERROGATIONS. 

(Cristispira  interrogationis,  Gross,  1910.) 
This  organism  was  also  found  by  Gross  in  Pecten  jaco- 
b&us.     It  is  25/4  in  length  by  0.5/1  in  breadth,  and  has 
pointed  ends,  bearing  apparently  to  Sp.  pectinis  a  rela- 
tion similar  to  that  of  Sp.  spiculifera  to  Sp.  anodontcs. 

SPIROCH^ETA  POLYSPIRA. 

(Wolff,  1907.) 

This  organism  was  found  by  Wolff  in  putrid  potato 
and  grows  on  ordinary  media.  He  describes  two 
forms :  one  with  sixty  to  seventy  curls,  70  to  8o/z  long 
by  1.5  to  i. 6/1  broad,  which  multiplies  by  longitudinal 
division,  and  a  second  140/1  long  by  0.25  to  0.3/1,  with 
sixty  to  eighty  curls.  Wolff  is  inclined  to  class  this 
organism  as  a  treponema.  It  is  placed  here  on  account 
of  its  length,  but  should  perhaps  rather  be  included  with 
the  small  spirochaetes. 

SMALL  SPIROCHJETES  OR  SPIRILLA. 

SPIROCH-ffiTA  OBERMEIERI. 

(Cohn,  1877.) 

(Sp.  recurrentis,  Lebert,  1874.) 

According  to  the  recent  descriptions  of  Novy  and 
Knapp  and  other  observers  this  spirochsete  or  spiril- 


SYSTEMATIC   DESCRIPTION   OF    SPECIES.  87 

him  is  from  7  to  19  p  in  length,  and  perhaps  £/*  in 
breadth  (Fig.  65) .  Long  forms  may  be  met  with,  up  to 
100 fj.  (i20fi,  Popovitch).  It  multiplies  by  transverse 
division  (see  Fig.  29),  usually  binary,  but  it  is  possible 
that  multiple  fission  may  at  times  occur.  It  usually 
stains  homogeneously  throughout,  with  the  exception 
that  the  ends  take  the  dye  rather  more  faintly.  Termi- 
nal appendages  or  flagella  may  be  present.  It  has  no 
undulating  membrane  or  lateral  flagella. 
Tangled  masses  of  spirochsetes  may  at  times 
be  seen,  or  two  or  more  individuals  twisted 
together.  Granules  are  occasionally  visible 
in  the  length  of  the  organism,  and  monili- 
form  degeneration  occurs  (see  Fig.  27).  Cox 
found  spirochsetes  inside  the  blood-corpuscles, 
where  they  broke  up  into  sections  and  finally 
into  granules. 

Norris,  Pappenheimer  and  Flournoy  state  spirochaeta 
that  they  cultivated  these  spirochsetes  in  (r|pU1Tober? 
citrated  blood.  The  spirochaetes  can  be  kept  meieri). 

i-         f  e     i  *.  -j     Ai      (Schellack.) 

alive  for  as  many  as  forty  days  outside  the 
body  (Novy  and  Knapp) ,  but  multiplication  under  such 
conditions  is  doubtful.  Destructive  (spirillicidal)  and 
agglutinating  bodies  are  formed  in  the  blood  of  im- 
munised animals,  and  passive  immunity  is  conferred 
by  injection  of  their  serum.  Apes,  rats  and  mice  can 
be  infected  with  these  organisms,  but  not  rabbits,  sheep, 
goats,  fowls  or  pigeons  (Fraenkel) . 

Fuchsine,  gentian  violet,  Giemsa's  stain  and  Levaditi's 
silver  method  may  be  used  for  staining  Sp.  obermeieri. 

It  is  said  that  infection  with  relapsing  fever  may  be 
carried  by  the  air.  The  bites  of  insects  have  also  been 
assigned  as  a  means  of  conveyance  of  the  disease. 
Klodnitsky  describes  a  development  of  the  spirochaetes 
into  long  filaments  within  the  bodies  of  bugs;  but 


88  SPIROCH^ETES. 

Nuttall  suggests  that  he  mistook  the  spermatozoa  of 

the  animal  for  spirochaetes  (see  Fig.  34). 

Sp.  obermeieri   is   destroyed   by   glycerine    (Gabrit- 

chewsky) .     The  action  of  a  variety  of  different  reagents 

upon  these  organisms  was  studied  and  tabulated  by 

MacKinnon. 

Sp.  obermeieri  in  some  form  or  other  is  capable  of 

passing  through  a  Berkfeld  filter. 

The    spirochaetes   met    with   in    cases    of  relapsing 

fever  in  America,  in  Bombay  and  in  African  tick  fever 
are  slightly  different  from  the  species 
met  with  in  Russian  relapsing  fever, 
the  typical  form  of  the  disease.  Sp. 
duttoni  of  tick  fever  is  described  be- 
low. The  spirochaete  of  American 
relapsing  fever  (Sp.  novyi)  is  more 
delicate  than  that  of  the  Russian 
disease  and  shows  more  regular  and 
closely-set  curls  (Fig.  66) .  The  spiril- 

lum  °f  Bomba7  fever  (SP-  carter$  is 

thinner,  less  regularly  curled,  and 
forms  loops  and  "figures  of  8"  (Novy  and  Knapp). 
Animals  inoculated  with  one  species  become  immune 
to  subsequent  infection  with  this  form,  but  not  to 
infection  with  the  other  species. 

SPIROCH^ETA  DUTTONI. 

(Novy  and  Knapp,  1906;  Breinl,  1906.) 

This  organism  was  first  discovered  by  Ross  and  Milne 
and  found  independently  by  Button  and  Todd.  It 
occurs  in  the  disease  known  as  tick  fever  on  the  east 
coast  of  Africa.  Infection  is  conveyed  by  the  bite  of 
the  tick,  Ornithodorus  moubata,  which  exists  in  great 
numbers  in  the  huts  of  the  natives.  The  spirochaete 
is  from  14  to  16/1  long,  with  six  to  seven  spiral  turns; 


SYSTEMATIC   DESCRIPTION   OF    SPECIES.  89 

long  forms  may  be  found  up  to  45/1  in  length.  It  has  a 
dark  central  rod  of  staining  material,  and  a  faint  sur- 
rounding periplast.  It  often  exhibits  dark  or  refrin- 
gent  granules  in  its  body.  Multiplication  is  probably 
by  transverse  division  (Fig.  67),  but  forked  forms  are 
found.  Rolled  up  specimens  are  seen  within  leucocytes, 
but  these  are  considered  by  Levaditi  and 
Mancunian  to  be  degenerative,  and  not 
resting  forms.  The  cilia  described  as 
present  on  these  spirochaetes  by  some 
writers  are  probably  artifacts  (Mayer). 
Koch  states  that  Sp.  duttoni  shows  no 
chromatin  granules,  but  Mayer  describes 
a  differentiation  of  nuclear  matter  and 
protoplasm  (see  Fig.  24).  The  latter 
writer  found  in  an  infected  mouse,  forms 
resembling  B.  fusiformis,  which  may  be 
stages  in  the  development  of  the  organ-  FlG  6  _Spiro. 
ism.  These  may  be  compared  with  chaeta  duttoni. 

^1  1        T^          j.   1  j   o-    j          (Schellack.) 

those  seen  by  Krzystalowicz  and  Sied- 

lecki  in  cases  of  syphilis  (possibly  stages  of  Sp.  pallidd) . 

L.  A.  and  R.  S.  Williams  state  that  they  cultivated 
Sp.  duttoni  in  defibrinated  blood  at  room  temperature, 
but  it  is  doubtful  whether  true  cultivation  was  effected. 
Duval  and  Todd  devised  a  special  medium  for  this  pur- 
pose (see  page  31)  and  succeeded  in  keeping  the  organ- 
isms alive  for  forty  days.  Levaditi  "cultivated" 
these  spirochaetes  in  collodion  sacs  in  the  peritoneal 
cavities  of  rabbits  along  with  the  serum  of  monkeys 
(Macacus  cynomolgus)  and  found  that  vibrio-forms 
developed. 

A  form  of  spirillum  fever  was  observed  by  Hodges 
and  Ross  in  Uganda,  which  is  believed  by  Moffat  to  be 
different  from  the  tick  fever  of  the  Zambesi  region. 
The  spirochastes  found  by  Hodges  and  Ross  were 


90  SPIROCHJETES. 

apparently  very  large  from  36  to  40^  in  length  by  4fi 
in  breadth. 

Allusion  has  already  been  made  (page  37)  to  the 
spirochaetes  found  by  Carter  in  S.  Arabia.  No  detailed 
description  of  the  organism  is  given,  but  the  writer 
describes  longitudinal  fission  as  taking  place  and  also  a 
process  of  conjugation.  Figures  40  and  48  are  copied 
from  his  original  paper  and  show  some  of  the  forms  ob- 
served ;  but  the  nature  of  their  appearance  is  so  peculiar 
that  it  would  be  premature  to  draw  definite  conclusions 
from  the  observations  thus  far  made. 

SPIROCH^ETA    PITHECI. 

(Thiroux  and  DufougerS,   1910.) 

Thiroux  and  Dufouger£  found  spirochaetes  in  mon- 
keys (Cercopiihecus  patas)  suffering  from  an  affection 
characterised  by  irregular  attacks  of  fever — a  condition 
resembling  relapsing  fever  in  man.  The  organisms 
were  15  to  20/x  in  length  by  o.  25/1  in  thickness,  and  pre- 
sented five  or  six  curls.  Rolled  up  forms  were  also  seen. 
The  blood  of  the  monkey  was  infective  between  the 
attacks  of  fever,  and  the  disease  was  readily  trans- 
missible to  mice,  less  easily  to  rats.  In  mice  the  incu- 
bation period  was  two  to  six  days,  and  death  usually 
occurred  on  the  twenty-fifth  or  twenty-sixth  day  after- 
ward. The  spirochaetes  were  present  in  one  case  in  the 
cerebro-spinal  fluid,  and  the  monkey  in  this  instance 
suffered  from  epileptiform  attacks.  The  authors  point 
out  a  resemblance  to  the  position  and  action  of  the 
trypanosomes  of  sleeping  sickness. 

SPIROCH^ETA    ANSERINA. 

(Sacharoff,  1890.) 

Sacharoff  found  these  organisms  in  geese  suffering 
from  a  fatal  malady,  characterised  by  fever,  wasting 


SYSTEMATIC   DESCRIPTION    OF    SPECIES.  QI 

and  diarrhoea.  The  bird  sits  apathetic  and  will  not 
eat.  The  joints  of  the  feet  are  so  tender  that  the  slight- 
est handling  of  them  causes  cries  of  pain.  The  disease 
is  a  septicaemia,  masses  of  spirochaetes  being  found  in 
the  blood.  According  to  Sacharoff  geese  and  ducks 
are  the  only  birds  susceptible,  the  incubation  period  of 
the  malady  being  two  days.  The  blood  of  the  infected 
bird  is  spirochaeticidal,  and  immunity 
is  produced  in  this  way,  not  by 
phagocytosis  (Gabritchewsky) . 
Sacharoff,  however,  saw  spirochaetes 
inside  leucocytes.  Gabritchewsky 
injected  a  horse  with  the  infected 
blood  and  obtained  a  preventive  and 

Curative  serum.  for   comparison. 

Cantacuzene  holds  that  bacterioly-    (Sacharo«-) 
sis  and  agglutination,  in  the  case  of  these  spirochaetes, 
are  phenomena  which  only  occur  in  vitro,  and  states 
that  destruction   of   the    organisms    takes    place    by 
phagocytosis  in  the  spleen. 

The  spirochaete  (Fig.  68)  is  from  10  to  30 /*  long  and 
thinner  than  Sp.  obermeieri.  It  multiplies  by  transverse 
division.  A  somewhat  curious  formation  is  depicted 
by  Cantacuzene  in  organisms  which  are  dividing,  viz., 
the  appearance  of  a  faint  rounded  nodule  in  the  centre 
at  the  point  of  division. 

Borrel  believes  that  Sp.  anserina  is  the  same  as  Sp. 
gallinarum,  but  this  is  not  consistent  with  SacharofFs 
statement  that  only  ducks  and  geese  are  susceptible  to 
the  former. 

SPIROCH-fETA     GALLINARUM. 

(Marchoux  and  Salimbeni,  1903.) 

According  to  Prowazek  these  spirochaetes  have 
flattened  bodies,  terminal  processes,  and  an  undulating 


Q2  SPIROCILETES. 

membrane  (?).  He  believes  that  they  possess  a  central 
rod  (axenstrang)  which  is  elastic,  and  a  contractile 
sheath.  In  length  they  vary  from  4  to  20;*,  and  may 
show  as  few  as  two  curls  or  a  large  number.  Terminal 
granules  are  often  seen,  and  the  chromatin  may  gather 
to  the  centre  of  the  organism,  leaving  the  ends  colour- 
less. Division  is  longitudinal.  Resting  forms  occur — 
some  with  terminal  loops,  others  closely 
curled  up,  others  again  in  the  form  of  a 
"figure  of  eight"  (see  Figs.  26,  33,  51). 
They  cling  together  in  masses,  but  no 
true  agglutination  occurs,  as  the  organ- 
isms free  themselves  again  from  the 
clumps.  They  may  get  inside  the  red 
blood-corpuscles  (Fig.  69).  Prowazek 

notes  that  the  undulating  membrane  is 
FIG.  69. — Spiro- 

chaeta  gailinarum  best  seen  in  macerated  specimens.  It 
de.  bl(P°rowazeko  is>  therefore,  probably  an  artifact.  The 
cilia  described  by  Zettnow  as  present 
on  these  organisms  are  also  probably  artifacts  and 
due  to  plasmolysis.  These  spirochaetes  are  killed  by 
glycerine. 

The  spirochaetes  can  be  directly  inoculated  from  fowl 
to  fowl,  and  are  also  conveyed  by  the  bites  of  ticks 
(Argas  miniatus,  A.  persicus,  A.  reflexus,  Ornithodorus 
moubatd) .  Rabbits  can  be  infected  with  this  organism 
(Levaditi  and  Lange) .  Marchoux  noted  a  loss  of  viru- 
lence in  these  spirochaetes  when  they  were  passed 
through  a  series  of  fowls  and  thinks  that  the  original 
virulence  is  regained  by  sojourn  in  the  body  of  a  tick. 

The  crisis  of  the  disease  is  not  due  to  the  formation 
of  antibodies,  but  may  be  due  to  phagocytosis.  Atoxyl 
is  both  protective  against  the  affection  and  curative 
when  it  already  exists.  Salvarsan  (Ehrlich-Hata)  is 
also  very  efficacious. 


SYSTEMATIC   DESCRIPTION   OF   SPECIES. 


93 


SPIROCH^TA     LAGOPODIS. 

(Fantham,   1910.) 

Fantham  found  this  spirochaete  in  the  blood  of  the 
grouse.  It  is  from  10  to  18/1  in  length,  and  relatively 
broad  (see  Fig.  25).  It  is  variable  in  form.  Its  ends 
taper  to  points,  and  it  has  an  undulating  membrane, 
spirally  wound  round  its  body.  Within  its  substance 
may  be  seen  a  series  of  chromatin  bars,  and  sometimes  a 
helicoid  core.  It  multiplies  by  both  longitudinal  and 
transverse  division.  It  may  be  transmitted  from  one 
bird  to  another  by  the  parasite,  Ixodes  ricinus. 

SPIROCH^TA    LOVATI. 

(Fantham,  1910.) 

This  organism  is  found  in  the  caecum  of  the  grouse. 
It  is  from  16  to  3  2  .  5/1  in  length,  and  its  ends  are  usually 
pointed,  rarely  rounded.  It  possesses  an  undulating 
membrane,  and  multiplies  by  both  transverse  and 
longitudinal  fission. 

SPIROCH^ETA  VESPERUGINIS. 

(Gonder,  1908.) 

Nicolle  and  Compte  found  spirilla 
in  a  Tunisian  bat  (Vesperugo  kuhlii). 
The  organisms  are  from  12  to  i8/* 
long,  and  £/*  or  less  in  breadth  ;  they 
have  pointed  ends  and  no  undulating 
membrane,  and  multiply  by  trans- 
verse division.  Crisis  and  relapse 
occur,  and  there  is  enlargement  of  the 
spleen;  some  immunity  is  conferred 
by  an  attack. 

Nicolle  and  Compte  regard  the  or- 
ganism as  a  bacterium,  but  Gonder 
claims  it  as  a  protozoon.  The  latter 
found  individuals  up  to  30^  in  length; 
and  small  forms,  3  to  5^  long,  were  x  2250.  (Gonder.) 


pIG.     7o.—  Spiro- 


94  SPIROCELETES. 

found  in  ticks  (Fig.  70) .  He  believes  that  this  spiro- 
chaete  divides  longitudinally  and  has  terminal  appen- 
dages of  periplast  and  an  undulating  membrane ;  but  as 
he  notes  that  this  last  only  becomes  visible  on  addition 
of  glycerine,  it  is  likely  that  the  apparent  membrane 
was  an  artifact.  Gonder  also  describes  dots  of  chro- 
matin  in  the  substance  of  the  organism,  which  at  other 
times  are  replaced  by  a  chromidial  network. 

SPIROCH^ETA  LUTR^E. 

(Prowazek,  1907.) 

This  organism  was  found  by  Prowazek  in  the  blood 
of  the  otter.  The  only  description  which  he  gives  of 
it  is  that  it  is  a  broad,  band-shaped  spirochsete,  with 
blunt  ends,  and  that  large  chromatin  bodies  (Chro- 


FIG.  71. — Spirochaeta  lutrae  and  rod-shaped  bodies.     (Prowazek.) 

matinbrocken)  were  visible  in  its  protoplasm  (Fig.  71). 
Whether  it  had  any  relation  to  certain  oval  bodies 
("like  p£brine  corpuscles")  present  in  the  blood  along 
with  it,  he  leaves  undetermined. 

SPIROCH.ETA  THEILERI. 

(Laveran,  1902.) 

This  organism  was  found  by  Theiler  in  cattle  suf- 
fering from  a  disease  characterised  by  fever,  diarrhoea, 
and  enlargement  of  the  spleen.  Disappearance  and  re- 
appearance of  the  spirochaetes  may  take  place,  as  in 
the  relapses  of  recurrent  fever.  The  parasite  is  from 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  95 

20  to  30/1  long  and  \  to  |/*  in  breadth;  it  appears 
in  regular  "cork-screws"  and  in  irregular  forms.  It 
shows  undulating  movements,  and  may  be  free  or 
attached  to  the  blood-corpuscles.  Infection  may  take 
place  by  the  bite  of  the  tick,  Rhipicephalus  decoloratus, 
and  possibly  a  developmental  stage  may  occur  in  this 
host,  as  direct  inoculation  did  not  succeed. 

Theiler  also  found  spirochaetes  in  horses  and  sheep 
suffering  from  fever,  and  suggests  that  these  may  be 
the  same  organisms.  He  succeeded  in  inoculating  a 
sheep  with  the  bovine  organism. 

Martin  found  spirochaetes  in  a  sick  horse  in  French 
Guiana;  this  form  measured  12  to  i$/y.  in  length  by 
\fj.  in  breadth. 

Baruchello  and  Pricolo  found  spirochastes  in  in- 
fective pleuropneumonia  of  horses;  these  organisms 
were  apparently  very  small,  measuring  3.2^  by  0.5^. 
They  occurred  in  the  spleen,  pulmonary  alveoli,  pleural 
effusion  and  blood.  They  sometimes  exhibited  terminal 
granules. 

Heanley  found  spirochaetes  in  buffaloes  in  China, 
and  Martoglio  and  Carpano  found  organisms  of  this 
nature  in  sheep.  The  relationship  between  all  these 
spirochaetes  is  doubtful.  Dodd  believes  that  the  spiro- 
chaetes met  with  in  the  horse,  ox,  and  sheep  (called 
by  Nov^y  and  Knapp  Sp.  equi,  Sp.  bovis,  and  Sp.  ovis, 
respectively)  are  identical. 

SPIROCH^TES  IN  THE  PIG. 

(Spirochaeta  suis?) 

Dodd  found  spirochaetes  in  a  disease  affecting  the 
skin  of  the  pig,  communicated  by  contact  and  inocu- 
lable  in  the  skin  of  a  healthy  animal.  The  organisms 
(Fig.  72)  were  not  found  in  the  blood;  they  disappeared 


96  SPIROCH^TES. 

and  reappeared  in  the  cutaneous  lesions  from  time  to 
time. 

Cleland  observed  spirochaetes  in  tumours  occurring 
at  the  site  of  castration  in  pigs.  The  organisms  were 
from  6  to  i2/x  in  length  and  exhibited  three  or  four 
irregular  curls.  Along  with  these  organisms  were  large 


FIG.   72. — Spirochaeta  suis.     (After  Dodd.) 

bacillary  forms,  some  of  which  were  slightly  undulat- 
ing and  suggestive  of  being  modified  forms  of  the 
spirochaetes. 

SPIROCELETA  LAVERANI. 

(Breinl  and  Kinghorn,  1906.) 

These  organisms,  found  in  mice  by  Breinl  and  King- 
horn,  measured  from  1.8  to  3.75^  in  length,  and  o.i 
to  0.2/4  in  breadth.  They  had  pointed  ends  (one  more 
so  than  the  other)  and  one  to  four  curls.  They  were 
transmissible  by  inoculation,  with  an  incubation 
period  of  five  days.  Transmission  by  fleas  and  lice 
was  not  effected.  The  mice  showed  enlargement  of 
the  spleen. 

SPIROCHAETA  MURIS. 

(Wenyon,  1906.) 

Found  by  Wenyon  in  mice,  this  organism  (Fig.  73), 
is  from  3  to  jn  long  and  2/1  broad.  It  has  no  undulat- 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  97 

ing  membrane,  and  multiplies  by  transverse  division. 
The  blood  of  the  mouse  is  not  infective,  nor  was  trans- 
mission by  fleas  effected.  It  is  apparently  a  harmless 
parasite. 

Wenyon  thinks  it  may  be  identical  with  the  form 
found  by  Borrel  in  mouse-cancer  and  perhaps  with  the 
spirilla  discovered  in  the  rat  by  Carter. 


FIG.  73. — Spirochaeta  muris.     (After  Wenyon.) 
SPIROCH^TA  MINOR. 

(Spirillum  minor,  Carter,  1887.) 

Carter  found  in  the  rat,  Mus  decumanus,  spirilla 
(spirochaetes  ?)  measuring  5  to  gft  in  length,  and  pos- 
sessing four  to  eight  curls.  He  describes  them  as 
shorter  and  slenderer  than  Sp.  obermeieri.  They  have 
pointed  ends,  and  exhibit  movements  of  rotation,  pro- 
pulsion, and  lashing.  They  disappear  and  reappear 
in  the  blood  of  the  rat,  being  replaced  during  their 
absence  by  granular  particles  and  filaments. 

SPIROCH^TES  IN  MOUSE-CANCER. 

Gaylord  found  spirochaetes  constantly  present  in 
mouse-cancers,  both  primary  and  secondary.  They 
were  studied  by  Calkins,  who  describes  them  as  2.5  to 
7.8/1  in  length,  o.6//  broad,  and  possessing  four  to 
thirteen  curls.  The  ends  are  blunt  and  rounded,  and 
they  have  no  undulating  membrane  or  flagella. 

Borrel  had  previously  observed  these  organisms  in 
1905.  They  are  not  causally  connected  with  the 
7 


98  SPIROCELETES. 

tumours,  but  are  passed  from  mouse  to  mouse  in  inocu- 
lating the  growths.  Calkins  identified  them  with  Sp. 
microgyrata  (see  below). 

SPIROCH^ETA  GONDII. 

(Nicolle,  1907.) 

Nicolle  found  spirochaetes  in  the  blood  of  the  rodent, 
Ctenodactylus  gondi.  They  measured  16  to  igji  in 
length  by  0.3^  in  breadth. 

SPIROCH^ETA  CULICIS. 

(Jaffd,  1907.) 

Jaffe  found  in  the  alimentary  canal  of  the  gnat  spiro- 
chaetes visible  with  a  comparatively  low  magnification 
(X  220).  They  were  flattened,  band-shaped  organisms, 
showing  deeply  stained  granules  in  their  protoplasm 
when  treated  with  Giemsa's  stain.  Their  movements 
were  lashing,  undulating,  and  "cork-screw-like."  They 
were  not  furnished  with  an  undulating  membrane,  and 
no  examples  of  division  were  seen. 

SPIROCH^TA  MINEI. 

(Prowazek,  1910.) 

These  organisms  are  found  in  the  stomachs  of 
"worker"  and  "soldier"  ants  (Termes lucifugus,  Rossi)  in 
Japan.  They  are  from  15  to  5o/x  in  length,  by  0.3  to  i/j. 
in  breadth.  The  extremities  are  pointed  and  chro- 
matin  granules  are  seen  in  the  protoplasm.  Division 
is  longitudinal  and  rolled  up  forms  and  rosettes  are 
found. 

SPIROCH^TA    GADI. 

(Neumann,    1909.) 

This  organism,  found  in  the  fish,  Gadus  minutus, 
resembles  Sp.  gallinarum,  measuring  10  to  i6/*  in  length, 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  99 

with  a  wave-length  of  3 . 5  to  4/z.  It  occurred  both  in 
the  peripheral  blood  and  in  the  internal  organs,  and 
was  actively  motile. 

SPIROCH^ETA    PELAMYDIS. 

(Neumann,   1909.) 

These  spirochaetes  were  found  in  the  blood  and  organs 
of  Pelamys  sarda.  They  are  shorter  and  thicker  than 
Sp.  gadi,  measuring  9  to  ioft  in  length,  occasionally  as 
much  as  18  to  20/1.  The  wave-length  of  the  curls  is  i 
to  1.9/1.  Movement  is  active. 

SPIROCH^TA  JONESII. 

(Button,  Todd,  and  Tobey,  1906). 

These  spirochaetes  from  the  fish,  Clavias  angoknsis, 
measured  i8/£  in  length  by  0.6/1  in  breadth.  They  ap- 
peared round  at  one  end  and  pointed  at  the  other,  a  fea- 
ure  which  suggests  transverse  division.  They  occurred 
both  isolated  and  in  clumps. 

SPIROCH^ETA  HARTMANNI. 

(Gonder,  1908.) 

This  small  spirochaete  was  found  by  Gonder  in  the 
intestine  of  Pinna  squamosa  and  P.  nobilis.  It  meas- 
ures 6  to  I4/*  in  length  by  i/i  in  breadth.  It  is  said 
by  Gonder  to  multiply  by  longitudinal  division,  on  the 
strength  of  certain  forked  forms  observed.  It  has 
pointed  ends  and  periplastic  appendages. 

SPIROCH.ETA    BUFONIS. 

(Dobell,  1908.) 

This  organism  was  found  by  Dobell  in  the  rectum 
of  the  toad  (Bufo  vulgaris) .  It  resembled  Sp.  buccalis, 
and  measured  8  to  io//  by  i.5/x.  It -had  an  undulating 
membrane.  The  same  writer  describes  another  form 


100  SPIROCILETES. 

in  the  large  intestine  of  the  toad,  which  he  compares 
with  Sp.  pallida  and  regards  as  a  treponema,  but  the 
illustrations  which  he  gives  do  not  show  the  same  regu- 
larity of  curls. 

SPIROCH^TA  BUCCALIS. 

(Cohn,  1877.) 

Several  spiral   organisms  are  found  in  the  human 
mouth.     The  largest  of  these,  Sp.  buccalis,  is  from  12 


FIG.   74. — Mouth  spirochaetes.     (Spirochaeta  buccalis.) 


FIG.  75.  FIG.  76. 

FIG.  75. — Spirochaeta  buccalis:  a,  c,  Showing  sheath;  b,  with 
terminal  filament;  a1,  Spirillum  sputigenum.  (Hoffmann  and 
Prowazek.) 

FIG.  76. — Spirochaeta  buccalis  (below).  Spirochaeta  dentium 
(above).  X  2200.  (Schaudinn.) 

to  20fi  in  length  and  £  to  i/i  broad  (Figs.  74,  75,  76). 
It  exhibits  long  undulations  (wave-length  2 p.,  Miihlens 
and  Hartmann)  rather  than  short  curls,  and  is  active  in 


SYSTEMATIC   DESCRIPTION    OF    SPECIES.  IOI 

its  movements.  Goadby,  however,  describes  spiro- 
chsetes  in  the  mouth,  apparently  Sp.  buccalis,  as  having 
a  sluggish  oscillating  movement.  It  has  terminal  peri- 
plastic  appendages,  and  Hoffmann  states  that  it  has 
an  undulating  membrane ;  this  is  doubtful.  Division 
is  said  to  be  longitudinal. 

SPIROCH^ETA  DENTIUM. 

(Koch,  1877.) 

This  small  spiral  organism  is  from  4  to  lo/t  long,  and 
exhibits  well-marked  regular  curls  (Figs.  76,  77,  78). 
Its  breadth  is  about  §/i.  It  very  closely  resembles  Sp. 
pallida  in  general  appearance,  but  is  said  to  be  rather 
thicker,  usually  shorter,  and  to  have  rather  less  sharply- 
wound  curls  (Muhlens  and  Hartmann) .  It  is  uncertain 


FIG.  77.  FIG.  78. 

FIG.  77. — Spirochaeta    dentium.        X   1500.       (Hoffmann    and 
Prowazek.) 

FIG.   78. — Spirochaeta  dentium.     Culture.     (Muhlens.) 

whether  these  organisms  have  any  pathogenic  influence ; 
they  are  found  chiefly  in  dirty  mouths,  in  the  tartar 
on  teeth,  and  in  the  discharges  of  pyorrhoea ;  and  Miller 
found  tangles  of  them  in  a  dental  abscess,  of  which  he 
thought  that  they  were  the  cause. 

These  organisms  may  perhaps  be  the  same  as  Loewen- 
thal's  Sp.  microgyrata  (see  below). 

An  intermediate  form,  between  Sp.  buccalis  and  Sp. 
dentium  is  also  described  (Sp.  media  (Prowazek);  (?) 
Sp.  denticola,  Arndt).  Spirillar  forms  with  lateral 
flagella  (Spirillum  sputigenum)  also  occur  (Fig.  75). 


102  SPIROCILETES. 

SPIROCH^JTA  VINCENTI. 

(Blanchard,  1906.) 

This  spirochaete  is  met  with  in  the  affection  known  as 
Vincent's  angina  (a  form  of  oral  and  pharyngeal  ulcer- 
ation  described  by  Vincent)  along  with  fusiform  bacilli. 
The  spirochastes  are  usually  about  lo//  long  (extreme, 
40 fi,  Mackie) ;  they  show  rather  sluggish  movements 
(see  Fig.  53).  Plaut  describes  an  euglenoid  movement, 
or  gradual  swelling  of  the  organism,  passing  like  a  wave 
along  its  length.  These  spirochaetes  are  constantly 
associated  with  fusiform  bacilli,  of  which  some  writers 
believe  them  to  be  a  developmental  form  (see  page  43). 
These  or  very  similar  associated  organisms  are  found 
not  only  in  the  throat,  but  also  in  gangrenous  processes 
in  other  parts  of  the  body.  The  spirochaetes  have  not 
been  satisfactorily  cultivated,  but  the  bacilli  may  be 
grown  on  artificial  media. 

The  resemblance  of  this  organism  to  Sp.  buccalis  is 
very  close,  and  some  writers  believe  that  they  are 
identical  (compare  also  Sp.  gracilis) . 


SPIROCH^ETA    GRACILIS. 

(Veszpr6mi,  1907.) 

This  organism,  found  by  Veszprdmi  in  a  case  of  ab- 
scess in  connexion  with  the  jaw,  in  a  man,  appears  to  be 
identical  with  Sp.  vincenti.  It  occurred  along  with 
fusiform  bacilli  and  cladothrix.  Levaditi  and  Stanesco, 
however,  obtained  this  organism  (?)  from  a  chancre  and 
cultivated  it  by  Schereschewsky's  method.  They  state 
that  it  is  a  distinct  species,  almost  indistinguishable 
from  Sp.  pallida :  it  exhibits,  however,  rather  less  regu- 
lar spirals,  is  rather  thicker,  moves  more  actively,  and  is 
stained  blue  by  Giemsa's  method.  The  description  is 
suggestive  of  Sp.  microgyrata  or  Sp.  pseudo- pallida. 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  103 

SPIROCH^ETA    SCHAUDINNI. 

(Prowazek,   1907.) 

Spirochaetes  and  fusiform  bacilli  are  found  in  the 
affection  known  as  tropical  ulcer  of  the  leg  (Fig.  79). 
Prowazek  states  that  they  are  band-shaped  organisms, 
possessing  an  undulating  membrane,  and  that  they 
multiply  by  longitudinal  fission.  In  a  specimen  (from 
case  of  this  disease?)  sent  to  me  by  Capt.  A.  Whitmore, 
from  Rangoon,  the  spirochaetes  were  accompanied  by  a 


FIG.  79.  FIG.  80. 

FIG.  79. — Spirochaete  of  ulcus  tropicum.  Sexual  forms.  X  2250. 
(Prowazek.) 

FIG.  80. — Spirochaetes  and  cladothrix  (?)  in  ulcus  tropicum. 
(From  a  specimen  by  Capt.  A.  Whitmore.) 

large  number  of  threads  resembling  filaments  of 
cladothrix  (Fig.  80).  The  spirochaetes  were  of  about 
the  same  thickness  as  the  threads,  and  stained  almost 
exactly  the  same  tint  with  fuchsine.  It  was  difficult  to 
resist  the  conclusion  that  they  were  forms  of  the  same 
organism.  (Vide  sub  Sp.  gracilis) . 

Spirochaetes  and  elongated  rod-shaped  bodies  were 
found  by  Assmy  in  phagedaenic  ulcers. 

SPIROCHAETES    IN    GANGRENOUS    PROCESSES. 

Polland  found  spirochaetes  in  cases  of  hospital  gan- 
grene; the  organisms  were  usually  from  15  to  20/4  in 
length,  but  forms  were  met  with  up  to  140;*.  They 


104  SPIROCILETES. 

usually  showed  five  or  six  blunt  curves,  and  occurred 
along  with  fusiform  bacilli. 

R6na  found  spirochaetes  in  noma,  hospital  gangrene, 
ulcus  gangrenosum  genitalium  and  pulmonary  gan- 
grene. They  occurred  along  with  fusiform  bacilli. 
Those  in  noma  and  gangrene  were  of  the  same  kind,  but 
those  seen  in  ulcus  gangrenosum  were  smaller. 

Possibly  the  latter  were  Sp.  microgyrata,  the  former 
Sp.  vincenti. 

SPIROCH^TA    PSEUDOPALLIDA. 

(Mulzer,  1905.) 

This  name  was  given  by  Mulzer  to  spirochaetes  closely 
resembling  Sp.  pallida,  but  not  identical  with  it,  found 
in  sores  on  the  genital  organs.  He  thinks  this  is  the 
same  organism  as  was  seen  by  Kiolomenoglou  and  Cube 
and  by  them  described  as  Sp.  pallida. 

SPIROCH^ETA    BRONCHIALIS. 

(Castellani,  1907.) 

Castellani  found  spirochaetes  in  cases  of  Haemorrhagic 
Bronchitis  occurring  in  Ceylon.  He  noted  the  presence 
of  four  different  lands:  (i)  a  thick  form,  with  irregular 
curls,  measuring  1 5  to  30^ ;  (2)  a  form  like  Sp.  refringens; 
(3)  a  thin  delicate  form  with  tapering  ends  and  small 
curls ;  and  (4)  another  thin  form  with  fewer  curls.  He 
speaks  of  an  acute  and  a  chronic  bronchial  spirochaetosis. 

Waters  also  found  spirochaetes  associated  with  febrile 
bronchitis,  and  Branch  noted  two  kinds  in  the  sputum 
of  a  tuberculous  patient  (along  with  tubercle  bacilli) — 
one  resembling  Sp.  pallida  and  the  other  like  Sp. 
refringens. 

Bertarelli  and  Volpius  found  spirochaetes  resembling 
Sp.  buccalis  and  others  resembling  Sp.  pallida,  but  stain- 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  105 

ing  more  readily,  in  the  expectoration  of  a  patient  suffer- 
ing from  heart-disease. 

SPIROCH^TA    MICROGYRATA. 

(Loewenthal,   1906.) 

Loewenthal   found    spirochaetes   in    the    discharges 
from  the  surfaces  of  ulcerated  cancers.     They  stained 
more  readily  than  Sp.  pallida.     Long  individuals  meas- 
ured 5  to  iifj.  by  i^  to  2//;  and  smaller 
specimens  2^  to  6/j.  by  &  to  £//.     The  small 
forms  showed  from  four  to  twelve  curls, 
with  a  wave-length  of  £//,  or  half  that  of 
Sp.   pallida.     Along  with  the  spirochaetes 
were  rod-shaped  and  bent  sausage-shaped      F.IG-  8l- 
bodies.      Figure    81   shows  one   of  these   0fPukerated 
organisms  as  depicted  by  Schaudinn. 

Krienitz  found  Sp.  microgyrata  in  cases 
of  cancer  of  the  stomach,  and  noted  that  the  form  of  the 
organism  changed  with  changing  conditions,  the  altera- 
tions involving  both  length,  thickness,  and  arrangement 
of  curls.  He  therefore  doubts  the  possibility  of  distin- 
guishing spirochaetes  by  their  morphological  characters 
alone. 

Calkins  identified  the  spirochaetes  found  by  Gaylord 
in  mouse-cancer  as  a  variety  of  Sp.  microgyrata  (see 
page  98). 

Hoffmann  found  spirochaetes  in  ulcerated  cancers 
along  with  fusiform  bacilli. 

SPIROCHAETES    IN     SMALL-POX. 

Sakurane  found  spirochaetes  in  the  lesions  of  a  patient 
suffering  from  small-pox,  both  in  the  contents  of  the 
pustules  and  in  the  tissues  below  these.  They  were 
long  and  slender,  and  showed  numerous  regular  curls; 
forms  with  irregular  curves  were  also  seen. 


106  SPIROCH^TES. 

SPIROCH^ETES    IN    VACCINE    LYMPH. 

Bonhoff  reported  the  presence  of  spirochaetes  in 
vaccine  lymph,  but  other  writers  believe  them  to  have 
been  artifacts. 

SPIROCHAETA    REFRINGENS. 

(Schaudinn,  1905.) 

This  spirochaete  was  found  by  Schaudinn  in  syphi- 
litic lesions  along  with  Sp.  pallida  (Fig.  82),  but  it  may 
occur  apart  from  the  latter  in  simple  lesions  of  the 
genital  regions.  It  is  broader  than  Sp.  pallida,  less 


FIG.  82.  FIG.  83. 

FIG.  82. — Spirochaeta  pallida  and  Sp.  refringens.  Smear  from 
a  syphilitic  chancre;  a,  Sp.  pallida;  b,  doubtful  forms;  c,  Sp. 
refringens. 

FIG.  83.     Spirochaeta  refringens.      (Schaudinn.) 

sharply  curled  (Fig.  83),  and  stains  more  readily  and 
deeply.  Its  length  is  from  8  to  12  p  and  its  breadth 
about  £/*.  It  has  terminal  periplastic  appendages,  but 
no  undulating  membrane  (Levaditi) .  Its  body  is  said 
to  be  flattened  (?),  and  not  round  like  that  of  Sp. 
pallida. 

Some  writers  believe  that  Sp.  refringens  is  the  same 
as  S^.  balanitidis  (Rille,  Kraus) .  It  may  be  the  same 
organism  as  was  described  by  Donn6  in  1837  as  Vibrio 
lineola  in  syphilitic  lesions.  Eitner  believes  that  more 


SYSTEMATIC   DESCRIPTION   OF   SPECIES. 


107 


than  one  species  of  spirochaete  is  included  under  the 
name  of  Sp.  refringens.  Richards  and  Hunt  distin- 
guished three  forms  of  Sp.  refringens,  somewhat  differing 
in  appearance,  and  also  believed  that  this  organism 
might  be  a  stage  in  the  development  of  Sp.  pallida  (see 
page  30). 

Sp.  refringens  was  cultivated  by  Levaditi  in  human 
blood  in  collodion  sacs  placed  in  the  peritoneal  cavities 
of  rabbits.  The  spirochaetes  developed,  showing  short 
vibrio-like  forms  along  with  long  spirochaetes ;  different 
species  of  bacteria  also  developed  in  symbiosis.  Inocu- 
lation of  these  cultures  failed  to  induce  balanoposthitis, 
and  Levaditi  does  not  think  the  spiro- 
chaetes  are  the  cause  of  this  disease. 

Sp.  refringens  was  found  by  Baer- 
mann  in  enlarged  glands  in  a  monkey 
infected  with  syphilis,  along  with  Sp. 
pallida;  it  may  therefore  become  a 
blood  parasite,  and  not  be  confined 
to  superficial  lesions. 

SPIROCH^TA    BALANITIDIS. 

(Hoffmann  and  Prowazek,  1906.) 
The  causal  (?)  organism  of  balanitis 
is  believed  by  Hoffmann  and  Prowazek 
to  be  distinct  from  Sp.  refringens; 
they  describe  it  as  a  band-shaped 
spirochaete  (Fig.  84)  with  six  to  ten 
curls,  \  to  f//  broad,  and  furnished 
with  periplastic  appendages  and  an 
undulating  membrane  (the  illustration  showing  the  later 
formation  is  not  convincing) .  It  exhibits  wave-like  and 
rotatory  movements.  It  is  stained  red  by  Giemsa's 
reagent.  Scherber  states  that  these  spirochaetes  can 
be  grown  anaerobically  on  media  containing  serum; 


FIG.  84. — Spiro- 
chaeta  balanitidis. 
a,  Terminal  fila- 
ments and  close 
curls;  b,  dividing; 
c,  supposed  undu- 
lating membrane. 
(Hoffmann  and 
Prowazek.) 


io8 


SPIROCH^TES. 


he  also  finds  that  they  make  their  way  into  the  blood- 
vessels and  are  not  confined  to  the  surface  of  the  ulcera- 
tive  lesions,  as  is  usually  supposed. 

SPIROCH^TA    PALLIDA. 

(Schaudinn,  1905.) 

(Spironema  pallida,  Vuillemin,  1905;  Treponema  pallidum, 

Schaudinn,   1905;  Spiroschaudinnia  pallida, 

Sambon,  1907.) 

This  organism  was  discovered  by  Schaudinn  in  March, 
1905,  and  announced  in  a  communication  by  Schaudinn 
and  Hoffmann.  It  is  usually  from  4  to  14^  long,  and 
exhibits  six  to  fourteen  curls;  but  longer  forms  are 
encountered,  up  to  40 /*  or  more.  Its  breadth  is  almost 
immeasurable,  and  may  be  £/*  or  less.1  It  has  pointed 


\ 


FIG.    85.— Spirochseta    pallida:    a,   After    Schaudinn;    b,    c,    after 
Krzystatowicz  and  Siedlecki. 

ends  and  terminal  appendages  of  periplast,  but  no 
undulating  membrane  (Fig.  85).  Its  body  is  round  in 
section,  and  the  curls  are  very  constant  and  regular.  It 
differs  from  most  other  spirochsetes  in  staining  a  red- 
dish colour  with  Giemsa's  stain,  instead  of  blue.  Its 
movements  are  rotatory  and  undulating,  occasionally 
lashing,  the  curls  remaining  constant  during  these 
motions.  In  certain  conditions,  however,  the  curves 

1  They  appear  considerably  thicker  when  stained  by  Levaditi's 
silver  method  than  with  ordinary  stains. 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  IOQ 

are  obliterated,  and  straight  and  other  atypical  forms 
are  seen  (Schaudinn).  Such  irregular  forms  are  com- 
mon in  "cultures"  (Levaditi  and  Macintosh,  Schere- 
schewsky) .  They  may  closely  resemble  Sp.  refringens. 
In  movement  the  long  slender  organ- 
isms may  become  short  and  plump 
(Krzystalowicz  and  Siedlecki). 
Forked  forms  of  Sp.  pallida  are  found 

and  also  specimens  showing  apparently 

.     ...    f,      ,     ,     •   ,     ,        '  T         ,      y       FIG.  86.— Spiro- 
two  individuals  twisted  together ;  from  chseta  pallida   in 

these  appearances  a  longitudinal  mode 
of  division  has  been  inferred,  but  Leva- 
diti states  that  division  is  transverse.  Refringent 
granules  may  be  seen  in  the  length  of  the  organism,  and 
also  terminally  placed. 

The  spirochaete  is  easily  found  by  mixing  a  loopful  of 
secretion  from  a  syphilitic  sore  with  a  drop  of  distilled 
water  on  a  slide,  dropping  on  a  cover-slip,  and  observing 


FIG.  87.     Spirochseta  pallida  in  leucocytes.     (Gierke.) 

in  the  "dark  field"  (see  p.  66).  Smears  from  such  a 
sore  may  be  fixed  in  osmic  vapour  and  stained  with 
Giemsa's  reagent,  or  fixed  and  stained  simultaneously 
with  this  fluid  (see  plate,  Fig.  A) . 

In  the  body  the  spirochastes  lie  chiefly  between  the 
cells  in  the  lymphatic  spaces,  in  lymphatic  vessels  and 
glands,  and  in  the  blood.  Occasionally  they  become 
intracellular  (Figs.  86,  87).  The  appearance  of  the 
organisms  in  the  liver  of  a  syphilitic  foetus,  when  stained 


110  SPIROCH^TES. 

by  Levaditi's  method,  is  shown  in  figure  88  (see  also 
plate,  Fig.  B). 

A  sexual  cycle  has  been  described  by  Krzystalowicz 
and  Siedlecki.  According  to  these  writers,  certain 
spirochaetes  become  thicker  and  less  sharply  curved, 
and  constitute  the  female  gametes.  Other  individuals 
are  found  with  many  nuclei,  and  these  by  fragmentation 
produce  a  large  number  of  minute  spirillar  forms, 
which  are  the  male  gametes  (Fig.  38).  One  of  these 
conjugates  with  a  female  gamete,  and  the  individual 


FIG.  88. — Spirochaeta  pallida  in  liver  of  syphilitic  foetus  stained 
by  Levaditi's  silver  method. 

thus  formed  probably  enters  on  a  resting  stage,  which 
has  not  been  definitely  identified.  These  writers  con- 
sider the  Spirochazta  pallida  to  be  a  trypanosome,  and 
call  it  Trypanosoma  luis.  The  same  view  is  taken  by 
Leuriaux  and  v.  Geets,  who  describe  oval  bodies  which 
elongate  and  give  rise  to  fine  filaments ;  these  latter  are 
the  microgametes  (the  spirochsetal  form),  while  the 
female  gametes  are  represented  by  a  tryponosome-like 
stage.  They  suggest  that  the  oval  forms  found  by 
them  are  identical  with  Siegel's  Cytoryctes  luis. 

The    observations   of   Krzystalowicz    and   Siedlecki 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  Ill 

have  not  been  confirmed.  Apart  from  the  sexual  cycle 
which  they  describe,  it  is  clear  that  if  the  variations 
in  the  form  of  Sp.  pallida  described  by  them  really 
occur,  the  foundations  of  the  whole  theory  of  its  causal 
connection  with  syphilis  are  shaken,  resting  as  they  do 
upon  the  morphological  constancy  of  this  organism,  with 
consequent  facility  of  identification,  and  its  occurrence 
only  in  syphilitic  lesions.  The  observations  of  these 
writers  would  point  to  the  existence  of  forms  indistin- 
guishable from  Sp.  rejringens  and  possibly  from  other 
organisms. 

Peculiar  bodies  were  found  by  Krzystalowicz  and 
Siedlecki  in  the  fluid  obtained  by  blistering  with  the 
actual  cautery  over  a  syphilitic  papule;  these  were 
much  larger  than  the  ordinary  spirochaetes  and  showed 
irregular  undulations  and  often  an  enlargement  at  one 
end.  They  were  irregular  in  shape  and  seemingly 
amoeboid.  Their  relationship  to  Sp.  pallida  was  un- 
certain. They  seem  to  be  the  same  bodies  as  were 
found  by  Horand,  who  noted  that  they  were  twenty- 
seven  or  twenty-eight  times  the  diameter  of  a  red  blood- 
corpuscle  and  had  a  head,  neck  and  tail.  Babes  and 
Panea  also  found  in  a  case  of  congenital  syphilis  bodies 
resembling  spermatozoa,  having  an  elongated  head,  i 
to  2/j.  long,  and  a  wavy  tail  or  two  tails.  They  were 
inclined  to  doubt  whether  these  were  parasites. 

Forms  somewhat  resembling  B.  fusiformis  were  noted 
by  Krzystalowicz  and  Siedlecki,  but  they  do  not  seem 
to  have  regarded  them  as  bacterial. 

Many  attempts  have  been  made  to  cultivate  Sp.  pal- 
lida. Recently  success  has  been  obtained  by  Schere- 
schewsky  and  by  Miihlens ;  and  Bruchner  and  Galacesco 
state  that  they  inoculated  rabbits  with  cultures  and 
produced  syphilis  (see  p.  33). 


112  SPIROCELETES. 

SPIROCH^ETA    PERTENUIS. 

(Castellani,  1905.) 
(Spirochaeta  pallidula,  Castellani,  1905.) 

Castellani  found  spirochaetes  present  in  eleven  out  of 
fourteen  cases  of  the  disease  called  yaws,  framboesia, 
or  pian;  and  the  discovery  was  confirmed  by  other 
observers  (Borne,  Schiiffner,  etc.).  The  organisms 
are  very  slender,  and  vary  in  length  from  a  few  microns 
up  to  1 8  or  20 p..  Their  extremities  are  pointed  as  a 
rule,  though  some  individuals  appear  to  have  blunt  ends. 
Their  curls  are  numerous  and  symmetrical,  but  some- 
times part  of  the  organism  may  appear  straight  (Fig. 
89).  Nodules  may  be  visible  at  their  extremities,  and 


FIG.  89. — Spirochaeta  pertenuis.     (Castellani.) 

also  in  their  length  (Fig.  21).  Ovoid  bodies  were  also 
found  by  Castellani  (5  to  8/t  by  4  to  6/*)  and  were 
thought  by  him  to  be  possibly  a  stage  in  the  develop- 
ment of  the  spirochaetes;  they  were  very  rare. 

Monkeys  were  inoculated  with  the  spirochaetes,  and 
the  organisms  were  found  in  the  resulting  lesions. 
Monkeys  thus  inoculated  showed  no  immunity  to 
syphilis;  this  is  held  to  prove  the  two  diseases  to  be 
different. 

These  spirochaetes  are  almost  exactly  like  Sp.  pallida, 
especially  when  stained  with  silver  by  Levaditi's 
method;  but  Mclntosh  states  that  the  positions  they 
respectively  occupy  in  the  lesions  are  different  (see  page 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  113 

57).     In  the  illustrations  given  by  Castellani  Sp.  per- 
tenuis  does  not  appear  so  regularly  curled  as  is  Sp. 

pallida. 

SPIROCH/ETA    OBTUSA. 
SPIROCH^ETA    ACUMINATA. 

(Castellani,  1905.) 

Along  with  Sp.  pertenuis  Castellani  found  other  forms 
of  spirochaetes  which  he  called  respectively  Sp.  obtusa — a 
thin  delicate  organism  with  blunt  extremities — and 
Sp.  acuminata,  also  very  slender,  but  with  tapering 
ends.  Wellman  could  not  identify  these  varieties  in 
the  lesions  of  yaws.  Their  nature  and  their  relation- 
ships to  Sp.  pertenuis  are  doubtful. 

SPIROCH^ETA  LYMPHATICA. 

(Proescher,  1909.) 

Proescher  and  White  found  spirochaetes  in  two  cases 
of  lymphadenoma ;  the  organisms  were  present  in 
enormous  numbers  in  the  .glands,  being  demonstrated 
by  Levaditi's  silver  method.  A  rhesus  monkey  was 
inoculated  with  material  from  the  glands,  and  developed 
first  a  local  nodule  at  the  site  of  inoculation,  and  sub- 
sequently further  secondary  nodules  in  distant  parts 
of  the  body.  Spirochaetes  were  found  in  all  the  lesions. 
A  second  monkey  was  successfully  inoculated  from  the 
first.  The  spirochaetes  resembled  Sp.  pallida  so  closely 
that  Proescher  at  first  believed  the  case  to  be  syphilitic. 1 

We  may  here  mention  a  spirochaete  found  by  Moritz 
in  a  case  of  severe  anaemia  and  cancerous  lymphangitis, 
the  organisms  being  present  in  the  bone-marrow  and  in 
the  wall  of  the  gut.  With  the  spirochaetes  in  the  latter 
position  were  bacterial  forms.  The  spirochaetes  were 

1  It  is  possible  that  there  was  actually  a  syphilitic  infection  in 
this  case. 


114  SPIROCTLETES. 

from  2  to  6/1  in  length,  and  had  from  three  to  ten  curls. 
They  differed  from  S^.  obermeieri  in  their  appearance, 
being  plumper  and  more  compact  (gedrungen) ;  they 
had  also  thicker  ends. 

SPIROCH^TA    ABORIGINALIS. 

(Cleland,  1909.) 

Spirochaetes  were  found  by  Wise  in  cases  of  the  de- 
structive local  lesion  called  ulcerative  granuloma  of 
the  pudenda  and  have  been  described  by  Cleland  and 


FIG.  90. — Sp.  aboriginalis  and  bacteria.      (Bosanquet.) 

by  Bosanquet  (Fig.  90).  The  organisms  are  about  12/1 
in  length,  ranging  from  forms  a  few  microns  only  in 
length  to  long  forms  attaining  18  or  20^.  They  are 
irregular  in  their  curls,  thus  being  clearly  distinguished 
from  Sp.  pallida,  and  from  Sp.  pertenuis,  as  seen  in  the 
tissues.  They  closely  resemble,  however,  the  pictures 
given  by  Castellani  of  Sp.  pertenuis  as  found  in  the 
secretion  of  yaws. 

The  spirochaetes  exist  in  the  tissues  along  with  bac- 
terial forms,  their  relationship  to  which  has  already 
been  discussed  (see  page  58). 

SPIROCH^TA   INTERROGANS. 

(Stimson,   1909.) 

Schaudinn  and  also  Novy  suggested  that  yellow  fever 
might  be  due  to  the  action  of  a  spirochaete,  but  failed  to 
find  such  organisms  in  cases  of  the  disease.  Stimson 
has  recently,  by  Levaditi's  method,  found  in  the  kid- 
neys of  one  such  patient  spirochaetes  (?)  which  often 


SYSTEMATIC   DESCRIPTION   OF   SPECIES.  115 

took  the  form  of  a  note  of  interrogation  and  which  he 
has  consequently  named  Sp.  interrogans.  They  are  14/1 
and  upward  in  length  by  about  £/*  in  thickness,  and  the 
wave-length  of  the  curls  is  i^  to  2fi.  They  were  situa- 
ted in  the  renal  cells  and  in  the  tubules,  but  not  in  the 
blood-vessels.  The  disco verv  has  not  been  confirmed. 


VARIOUS  DOUBTFUL  SPIROCH^ETES  AND  SPIRILLA. 

Adele  Oppenheimer  found  spirochsetes  in  the  mucus 
of  the  alimentary  canal  of  the  dog;  some  exhibited 
snake-like  movements  (lashing  or  wriggling),  while 
others  had  only  cork-screw  movements,  the  curls  in 
the  body  remaining  fixed. 

These  may  correspond  with  the  spirilla  discovered 
by  Bizzozero  in  the  dog ;  these  organisms  had  three  to 
seven  curls,  and  were  3  to  Sft  long;  they  lay  within 
vacuoles  in  the  epithelial  cells.  Salomon  also  found 
spirilla  in  the  intestines  of  dogs,  cats  and  rats;  these 
were  longer,  and  he  distinguished  three  forms — a  thick 
form  with  seven  to  nine  curls,  thicker  at  the  middle 
than  at  the  ends;  a  long  form  with  fifteen  to  twenty- 
four  curls,  the  axis  being  often  bent ;  and  a  form  about 
as  long  as  the  first,  but  with  only  two  to  five  wavy 
curls.  They  had  terminal  flagella  and  lay  in  the  mucus 
of  the  alimentary  canal  and  also  in  vacuoles  in  the  cells. 
Rigaud  found  spirilla  like  Spirochata  pallida  in  the 
stomachs  of  dogs  and  cats,  lying  at  the  bottom  of  the 
peptic  glands.1 

Spirilla  were  found  by  Kowalski  in  the  dejecta  of 
cholera  patients,  resembling  Sp.  dentium.  They  had 
two  to  three  curls  and  pointed  ends,  and  did  not  grow 
in  culture  media  (Abel) .  Rechtsamer  found  that  they 

lCf.  Balfour's  discovery  of  spirochaetes  in  ulcers  of  the  intestines 
of  dogs  and  monkeys,  recorded  on  page  44. 


Il6  SPIROCH^ETES. 

lived  a  little  while  in  broth :  he  also  noted  that  the  more 
cholera-vibrios  there  were  present,  the  fewer  were 
the  spirilla,  and  vice  versa. 

Le  Dantec  found  spirilla  in  the  stools  of  certain  pa- 
tients suffering  from  dysentery. 

Werner  found  in  the  human  alimentary  canal  two 
forms  of  spirochaetes — one  with  long  undulations,  4 . 6 
to  7 . 3/1  in  length,  which  may  occur  in  masses  (this  form 
is  not  identical  with  any  of  the  mouth-spirochsetes) ;  and 
a  second,  with  finer  curls,  3.5  to  6 .  i/i  in  length,  resem- 
bling Sp.  dentium.  The  former  he  calls  Sp.  eurogyrata 
and  the  latter,  Sp.  stenogyrata. 

Smith  (Theobald)  found  spirilla  along  with  comma 
bacilli  (vibrios)  in  the  intestines  of  pigs ;  they  had  two 
to  three  curls,  with  a  wave-length  of  2//.  He  could 
not  cultivate  them. 

Novy  and  Knapp  found  spirilla  (Spirillum  glossince) 
in  the  stomach  of  the  tse-tse  fly;  short  forms  measured 
8/z  in  length  and  long  ones  1 5^.  The  spirilla  were  shorter, 
narrower,  and  more  closely  curled  than  Sp.  obermeieri. 
They  multiplied  by  transverse  division,  the  long 
forms  probably  being  about  to  divide  (see  also  page 
68). 

Mezinescu  found  spirilla  in  the  pus  from  a  case  of 
calculous  pyelitis ;  they  were  from  3.6  to  8//  long,  rarely 
attaining  a  length  of  10  or  i2/x.  Doerr  found  spirilla 
in  the  pleural  and  pericardial  fluid  of  a  syphilitic  sub- 
ject and  thinks  they  may  have  been  the  same  as  those 
described  by  Mezinescu. 


BIBLIOGRAPHY. 

(*)Papers  thus  marked  I  have  not  been  able  to  consult  in 
the  original. 

GENERAL,    CLASSIFICATION,    ETC. 

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Blanchard.     Semaine  medicate,  1906,  I,  i. 

Gonder.     Centralbl.f.  Bakt.,  1909,  XLIX,  190. 

Gross,     Mitih.  a.  d.  Zool.  Station  zu  Napel,  1910,  XX,  41. 

Leishman.     Journ.   Prevent.    Medicine,    1906,    XIV,    513. 

Levaditi  and  Rosenbaum.     Ibid.,  1908. 

Miihlens.     Zeitschr.f.  Hygiene,  1907,  LVII,  405. 

Novy.*     Proc.  Pathol.  Soc.  of  Philadelphia,  1907. 

Prowazek.     Centralbl.f.  Bakt.,  1908,  XL VI,  229. 

Reichert.*     Hygien.  Rundschau,   1907,  XVII,   1121. 

Schwellengrebel.  Compt.  rend.  Socitte  de  Biologie,  1907, 
LXII,  213. 

Schwellengrebel.  Annales  de  I'Inst.  Pasteur,  1907,  XXI, 
448,  562. 

Schwellengrebel,     Centralbl.f.  Bakt.,  1909,  XLIX,  529. 

Schilling.     Centralbl.  f.  Bakt.,   1908,  Suppl.  to  XLII,  80. 

STAINING    METHODS. 

Bertarelli  and  Volpino.     Centralbl.  f.  Bakt.,  1906,  XLI,  74. 

Davidsohn.     Berlin,  klin.  Wochenschr.,   1905,  XLII,  985. 

Foix  and  Mallein.     C.  R.  Soc.  de  Biol.,  1907,  LXII,  1201. 

Forest.     Centralbl.f.  Bakt.,  1906,  XLII,  608. 

Giemsa.     Deutsch.  med.   Wochenschr.,   1905,  XXXI,  1026. 

Goldhorn.*  Postgraduate,  1906,  No.  2  (abstr.  Zentralbl. 
f.  inn.  Med.,  1906,  707). 

Herxhiemer  and  Hubner.  Deutsch.  med.  Wochenschr., 
1905,  XXXI,  1023. 

Hoffmann  and  Beer.     Ibid.,  1906,  XXXII,  869. 

Levaditi.     Ann.  de  I'Inst.  Pasteur,  1906,  XX,  41. 

Levaditi.     C.  R.  Soc.  de  Biol.,  1906,  LX,  67. 

MacNeal.  Journ.  Amer.  Med.  Assoc.,  1907,  XL VIII,  609. 
117 


Mandelbaum.     Muenchener  med.  Wochenschr.,  1907,  LIV, 
2268. 

Meirowsky.     Ibid.,  1910,  LVII,  777. 

Oppenheimer    and    Sachs.     Deutsch.    med.     Wochenschr., 
1905,  XXXI,  1156. 

Petresco.     Ibid.,  1905,  LIX,  680. 

Proca  and  Vasilescu.     C.  R.  Soc.  de  Biol.,  1905,  June  29. 

Proca    and    Vasilescu.*     Rivista    Stiintelor.    Med.,  1905, 
2  (abstr.  Zentralbl.f.  -inn.  Med.,  1906,  131). 

Proescher  and  White.     Journ.  Amer.  Med.  Assoc.,  1908, 
XLIX,  1908. 

Ravaut  and  Ponselle.    C.  R.  Soc.  de.  BioL,  1908,  LXV,  438. 

Reitmann.     Deutsch.  med.  Wochenschr.,  1905,  XXXI,  997. 

Sabraz^s  and  Duperrie\     C.  R.  Soc.de  Biol.,  1909,  LXVI, 
690. 

Schereschewsky.  Centralbl.  f.  Bakt.,  1907,  XLV,  91. 

Schmorl.     Deutsch.  med.  Wochenschr.,  1907,  270. 

Simonelli  and  Bandi.     Centralbl.  f.  Bakt.,  1906,  XL,  159. 

Von  dem   Borne.     Joiirn.    Tropical  Med.,  1907,   X,    345. 

Yamamoto.*     Centralbl. f.  allg.  Path.  u.  path.  Anat.,  1909, 
Feb.  27. 

Yamamoto.     Centralbl.  f.  Bakt.,  1908,  XLVII,  571. 

DARK  FIELD  EXAMINATION. 

Bayly.     Practitioner,  Feb.,  1910. 

Gastou  and  Commandon.     Bull.  Soc.  med.  des  Hdpitaux, 
Mar.,  1909. 

Harrison.    Journ.  R.  Army  Med.  Corps,  1910,  XIV,  506. 
Ogilvy.     Lancet,  Oct.  2,  1909. 

INDIAN-INK  METHOD. 

Burri,     Das  Tiischeverfahren,  Jena,  1909. 
Fruhwald.     Berlin,  klin.  Wochenschr.,  1909,  2121. 
Gins.     Centralbl.  f.  Bakt.,  1919,  LII,  620. 
Hecht  and   Wilenko.      Wiener  klin.    Wochenschr.,    1909, 
XXII,  932. 

Plaut.     Berlin,  klin.  Wochenschr.,  1909,  2276. 
Rosenhauer.     Muench.  med.  Wochenschr.,  1909,  2394. 

PSEUDO-SPIROCH^TES. 

Beitzke.     Centralbl.  f.  Bakt.,  1907,  XLIII,  369. 
Friedenthal.     Berlin,  klin.  Wochenschr.,  1907,  99. 


BIBLIOGRAPHY.  1 1 9 

Gierke.     Ibid.,  Jan.  21. 

Meyer.     Centralbl.f.  Bakt.,  1908,  XLVL  319,  339. 

Omeltschenko.*  Rousski  Vratch.,  July  23,  1905  (abstr. 
New  York  Med.  Journ.,  Sept.  30,  1905). 

Baling.     Centralbl.  f.  Bakt.,  1907,  70,  162,  229,  362. 

Saling.     Centralbl.f.  Bakt.,  1906,  XLII,  38,  120. 

Schulze.  Berlin,  klin.  Wochenschr.,  1906,  No.  37,  XLIII, 
1213. 

Schulze.*     Klin.  Monatshefte  f.  Augenheilk.,  1907,  XLV. 

Wolff.     Ibid.,  1907,  XLIII,  156,  222. 


SP.    PLICATILIS. 

Biitschli.*  Ueber  dem  Bau  der  Bakt.  u.  verwandten  or- 
ganismen,  Leipsig,  1892. 

Biitschli.     Deutsch,     med.     Wochenschr.,     1906,     No.     2. 

Doflein.     Lehrb.  der  Protozoenkunde,  1909. 

Ehrenberg.*     "Die   Infusionsthierchen,"   Leipsig,  1833. 

Laptschinsky.*  Centralbl.  f.  d.  med.  Wissensch.,  1880 
XVIII,  319. 

Schaudinn.  Arb.  a.  d.  kaiserl.  Gesundheitsamte,  1907, 
XXVI,  u. 

Schaudinn.     Deut.  med.  Wochenschr.,  1905,  No.  42. 

Zuelzer.     Zoologishe  Anzeiger,  1910,  XXXV,  795. 


SP.    BALBIANII. 

Borrel  and   Cernovodeanu.     C   R.   Soc.    de  Biol.,    1907, 
XLII,  1102. 

Certes.*     Bull  Soc.    Zool.    de    France,    1882,    VII,    347. 
Fantham.  Ann.  and  Mag.  of  Natural  History,  1907,  XIX, 

439- 

Fantham.    Quart.  Journ.    of  Micr.  Sci.,  1908,  LII,  pt.   1. 
Fantham.     Parasitology,  1910,  II,  392. 
Rolling.     Centralbl.  /.  Bakt.,  1907,  XLIV,  665. 
Laveran  and  Mesnil.     C.  R.  Soc.  de  Biol.  1901,881. 
Lustrac.*     Act.  Soc.  Linn,    de   Bordeaux,    1896,   X.   265. 
Perrin.     Arch.  f.  Protistenkunde.,  1906,  VII,  131. 
Schwellengrebel.     C.  R.  Soc.  de  Biol.,    1907,  LXII,  214. 
Schwellengrebel.     Centralbl.    f.    Bakt.,    1908,    XLVI,    i. 
Vtes.     C.  R.  Soc.  de  Biol.,  1906,  LXI,  408. 


120  SPIROCILETES. 

SP.    ANODONT.E. 

Keysselitz.  Arb.  a.  d.  kaiserl.  Gesunheitsamte,  1906, 
XXIII,  566. 

Schellack.     Ibid.,  1909,  XXX,  379. 

SP.    PINNAE. 

Gonder.  Centralbl.  f.  Bakt.,  1908,  XLVII,  491;  1909, 
XLIX,  191. 

SP.    MACTR.E. 

Prowazek.  Arch.  /.  Schiffs.  u  .Tropenhygiene,  1910,  XIV, 
297. 

SP.    PECTINIS.       SP.    INTERROGATIONIS. 

Gross.     Mitth.  a.  d.  Zool.  Station  zu  Napel,  1910,  XX,  41. 

SP.    POLYSPIRA. 

Wolff.     Centralbl.  f.  Bakt.,  1907,  XVIII,  Ref.,  448. 

SP.OBERMEIERI.1 

Bardach.     Ann.  de  I'lnst.    Pasteur,  1899,  XIII,    365. 

Bertarelli.     Centralbl.  f.  Bakt.,  1906,  XLI,  492. 

Bohne.     Arch.  f.  Schiffs.  u.  Tropenhygiene,  1908,  XII,  343. 

Browse.     British  Med.  Journ.,  1905,  I,  532. 

Carlisle.     Journ.  Infect.  Diseases,  1906,  III,  233. 

Cohn.*     "  Beitr.  z.  Biol.  der  Pflanzen,"  1877. 

Cox.     Brit.  Med.  Journ.,  1906,  I,  1400. 

Fischer.     Centralbl.  f.  Bakt.,  1907,  XL VI,  715. 

Fraenkel.     Berlin,  klin.  Wochenschr.,  1907,  V,  68 1. 

Fraenkel.     Centralbl.  f.  Bakt.,  1908,  XLVII,  349,  471. 

Fiilleborn  and  Mayer.     Mediz.  klinik.,  1907,  III,  487. 

Gabritchewsky.     Ann.  de  I'lnst.  Pasteur,    1896,   X,   630. 

Gabritchewsky.  Centralbl.  f.  Bakt.,  1906,  XXXVII, 
Ref.,  397. 

Karwacki  and  Szokalski.  C.  R.  Soc.  de  Biol.,  1910,  CXVIII, 
228,  286,  449. 

Klodnitzky.     Ibid.,  1907,  XLV,  126. 

Kretz.  Wien.  klin.  Wochenschr.,  1906,  XIX,  1462. 

Lafforgue.     Semaine  medic ale,  1905,  152. 

1  Only  recent  papers  are  quoted. 


BIBLIOGRAPHY.  121 

Mackinnon.     Parasitology,  1907,  II,  267,  281. 

Manteuffel.  Arbeiten  a.  d.  kaiserlichem  Gesundheitsamte , 
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Oppenheimer.*  Collected  Studies  from  the  Research  Lab., 
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Popovitch.      Th^se  de  Paris,  1906,  No.  158. 

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Schellack.     Ibid.,  1909,  XXX,  359. 

SergentandFoley.*    Bull.  Soc.  Path.  Exotique,  1908,  I,  174. 

Soudakewitch.     Ann.   de  I'lnst.   Pasteur,    1891,    V,    515. 

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SP.    DUTTONI. 

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I.I57 

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122  SPIROCBLETES. 

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Koch.     Berlin,  klin.  Woch.,  1906,  XLIII,  185, 

Leishman.  Trans.  Soc.  Trop.  Med.  and  Hygiene,  Jan., 
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619,  815. 

Manteuffel.  Arb  a.  d.  kaiserl  Gesundheitsamte,  1908, 
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Mayer.*     Arch.  f.  Schiffs-  u.  Tropen-hygiene,  1908,  XII,  7. 

Moffat.     Lancet,  1907,  Jan.  26. 

Newstead.     Liverpool  Sch.   of  Trop.   Med.,   Mem.  XVII. 

Ross  and  Milne.     Brit,  Med.  Journ.,  1904,  II,  1453. 

Stephens.     Lancet.  1906,  II,  438. 

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Tedeschi.     Centralbl.  f.  Bakt.,  1910,  LIV,  12. 

Vassal.     C.  R.  Soc.  de  Biol.,  1907,  LXII,  414. 

Williams,  L.  and  S.  Liverpool  Sch.  of  Trop,  Med.,  Mem. 
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SPIROCH^TA    PITHECI. 

Thiroux  and  Dufougere".  Compt.  rend.  Acad.  des  Sciences, 
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SP.    ANSERINA. 

Borrel.     C.  R.  Soc.  de  Biol.,  1906,  LX,  138. 
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439.  635.  721,  778. 
Sacharoff.     Ann.  de  I'lnst.  Pasteur,  1891,  V,  564. 

SP.  GALLINARUM. 

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Balfour.  Journ.  Trop.  Med.,  1909,  X,  I. 

Bevan.*  Journ.  Comp.  Path,  and  Therap.,  1908,  XXI, 
i,  43- 


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SP.    LAGOPODIS.      SP.    LOVATI. 

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SP.    VESPERUGINIS. 

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XX,  311. 


124  SPIROCILETES. 

SP.    LUTR.E. 

Prowazek.  Arb.  a.  d.  kaiserl.  Gesundheitsamte,  1907, 
XXVI,  31. 

SP.    THEILERI. 

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Laveran  and  Vall£e.  C.  R.  Acad.  des  Sciences,  1905, 
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SP.    SUIS. 

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Cleland.     Parasitology,  1908,  218. 

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SP.    LAVERANI.       SP.    MURIS.       SP.      MINOR. 

Breinl  and  Kinghorn.     Lancet,  1906,  II,  Sept.  8. 

Carter.     Sci.  Mem.  by  Officers  of  the  Indian  Army,  1887, 

HI,  45- 

Wenyon.      Journ.  of  Hygiene,  1906,  VI,  580. 

SP.    IN    MOUSE    CANCER. 

Calkins.     Journ.  Infect.  Dis.,  1907,  IV,  171. 
Borrel.     C.  R.  Soc.  de  Biol.,  1905,  LVIII,  780. 
Deetjen.     Muench.  med.  Wochenschr.,  1908,  No.  22. 
Gaylord.    Journ.  Infect.  Dis.,  1907,  IV,  155. 
Gaylord.     Berlin,  klin.  Wochenschr.,  1908,  2296. 

SPIROCH^ETA    GONDII. 

Nicolle.     Compt.  rend.  Soc.  de  Biologie,  1907,  LXIII,  213. 


BIBLIOGRAPHY.  125 

SP.   CULICIS. 

Jaffe.     Arch.f.  Protistenk.,  1907,  IX,  100. 

SP.    MINEI. 

Prowazek.    Arch.    f.    Schiffs.    u.    Tropen-Hygiene ,     1910, 

XIV,  297. 

SPIROCH^TA    GADI.       SP.    PELAMYDIS. 

Neumann.     Zeitschr.f.  Hygiene,  1909,  LXIV,  i. 

SPIROCH^TA    JONESII. 

Button,  Todd  and  Tobey.     Journ.  of  Med.  Research,  1906, 

XV,  491. 

SP.   HARTMANNI. 

Gonder.  Centralbl.  f.  Bakt.,  1908,  XL VII,  491;  1909, 
XLIX,  19 1. 

SP.    BUFONIS. 

Dobell.     Quart.  Journ.  Micr.  Sci.,  1908,  LII,  121. 

SP.    BUCCALIS.       SP.    DENTIUM. 

Arthaud.     C.  R.  Acad.  des  Sciences,  1909,  CXLIX,  1409. 
Bertarelli  and  Volpino.    Centralbl.  f.  Bakt.,  1906,  XXXVII, 

»i«. 

Feldmann.     Wiener  klin.  Wochenschr.,  1906,  XIX,  695. 

Goadby.     Lancet,  1906,  II,  1281. 

Hoffmann.  Centralbl.  f.  Bakt.,  1906,  XXXVIII,  Ref., 
Beiheft,  112. 

Hoffmann  and  Prowazek.     Ibid,,  1906,  XLI,  818. 

Miller.     Deutsch.    med.    Wochenschr.,    1906,  XXXII,    348. 

Schaudinn.  Arb.  a.  d.  kaiserl.  Gesundheitsamte,  1907. 
XXVI,  ii. 

Zettnow.     Zeitsch.f.  Hygiene,  1906,  LII,  485,  539. 

SP.    VINCENTI.1 

Eichmeyer.     Jahrb.  f.  Kinderheilk.,   1905,  XIII,  65. 
Ellermann.     Zeitschr.f.  Hygiene,  1907,  LVI,  453. 

'For  list  of  other  articles  see  Beitzke:  Centralbl.  f.  Bakt.,  1904, 
XXXV,  orig. 


126  SPIROCH^ETES. 

Launois  and  Loederich.     Bull.  Soc.  m6d.  des   Hdspitaux, 

1905,  XXII,  601. 

Lewkowitz.     Centralbl,f.  Bakt.,  1906,  XLI,  153. 

Mackie.     Lancet,  1905,  II,  no. 

Miihlens  and  Hartmann.  Zeitschr.  f.  Hygiene,  1906, 
LV,  81. 

Plaut.     Centralbl.f.  Bakt.,  1907,  XLIV,  310. 

Popovitch.      Thlse  de  Paris,  1906,  No.  158. 

Reiche.     Muench.  med.  Wochenschr.,  1905,  No.  32. 

Scherber.     Dermatol.  Zeitschr.,  1907,  XIV,  88. 

Schmiedlechner.  Zeitschr.  f.  Gebiirtshilfe  u.  Gyn.,  1905, 
LVI,  291. 

Tunnicliffe.    Journ.  Infect.  Dis.,  1906,  148. 

Vincent.     Lancet,  1905,  I,  1260. 

Vincent.     Muench.    med.    Wochenschr.,    1905,    LII,    1287. 

Weaver  and  Tunnicliffe.    Journ.  Infect.  Dis.,  1905,  II,  446. 

Weaver    and    Tunnicliffe.     Journ.    Amer.    Med.    Assoc., 

1906,  XL VI,  481. 

Wright.     Lancet,  1904,  II,  73. 

SP.  GRACILIS. 

Levaditi  and  Stanesco.  C.  R.  Soc.  de  BioL,  1909,  LXVII, 
188. 

Veszprdmi.  Centralbl.  f.  Bakt.,  1907,  XLIV,  408,  515, 
S32-  648. 

SP.  SCHAUDINNI. 

Assmy.     Arch.  f.  Schiffs.  u.  Tropenhyg.,  1909,  XIII,  p.  657. 
Keysselitz  and  Mayer.     Ibid.,  1909,  XIII,  137. 
Prowazek.     Arb.    a.    d.    kaiserl.    Gesundheitsamte,    1907, 
XXVI,  23. 

SP.    IN    GANGRENOUS    PROCESSES. 

Polland.     Wien.  klin.  Wochenschr.,  1905,  XVIII,  1236. 

Prowazek.  Arb.  a.  d.  kaiserl.  Gesundheitsamte,  1907, 
XXVI,  23. 

R6na.  Verhandl.  d.  deutsch.  dermatol.  Ges.  (1906),  1907, 
IX,  471- 

SP.    PSEUDO-PALLIDA. 

Mulzer.     Berlin,  klin.  Wochenschr.,  1905,  XLII,  1144. 


BIBLIOGRAPHY.  127 

SP.    IN    BRONCHIAL    SECRETION. 

Bertarelli  and  Volpino.     Centralbl.f.  Bakt.,  1906,  XXXVII, 
Ref.,  230. 

Branch.     Brit.  Med.  Journ.,  1906,  II,  1537. 
Castellani.     Lancet,  1906,  May  19. 
Castellani.    Journ.  Trop.  Med.,  1909,  399. 
Waters.     Lancet,  1909,  I,  689. 

SP.    MICROGYRATA. 

Hoffmann.     Berlin,  klin.  Wochenschr.,  1905,  880. 
Krienitz.     Deutsch.  med.  Wochenschr.,  1906,  XXXII,  872. 
Krienitz.     Centralbl.f.  Bakt.,  1906,  XLII,  43. 
Loewenthal.     Berlin,  klin.  Wochenschr.,  1906,  XLIII,  283. 
Simmonds.     Verhandl.  d.  deutsch.  pathol.  Ges.,  1908,  116. 

SP.    IN    SMALL-POX. 

Sakurane.*     Vortrag  in  d.  med.  Gesellsch.  z.  Osaka,  May 
20,  1907  (abstr.  Centralbl.f.  inn.  Med.,  1907,  835). 

SP.    IN    VACCINE-LYMPH. 

Bonhoff.     Berlin,   klin.    Wochenschr.,    1905,    XLII,    1142. 
Carini.     Centralbl.f.  Bakt.,  1905,  XXXIX,  685. 
Miihlens  and  Hartmann.     Ibid.,  1906,  XLI,  49. 
Supfle.     Ibid.,  1906,  XL,  495. 

SP.    REFRINGENS. 

Dreyer.     Deutsch.  med.  Wochenschr.,  1907,  No.  18. 
Eitner.     Muench.  med.  Wochenchr.,  1907,  April  16. 
Levaditi.     C.  R.  Soc.  de  Biol.,  1906,  LXI,  182,  185. 
Richards  and  Hunt.     Lancet,  1906,  I,  667. 
Rille.     Muench.  med.  Wochenschr.,  1905,  1377. 
Schaudinn.     Arb.   a.    d.   kaiserl.     Gesundheitsamte,     1907, 
XXVI,  ii. 

Schaudinn  and  Hoffmann.     Ibid.,   1905. 

SP.    BALANITIDIS, 

Hoffmann  and  Prowazek.     Centralbl.  f.  Bakt.,  1906,  XLI. 
741. 

Kraus.     Arch.  f.  Dermatol.  u.  Syph.,  1906,  LXXX,  255. 


128  SPIROCILETES. 

SP.   PALLIDA. 

Alvarez.     Journ.  Amer.  Med.  Assoc.,  1906,  XL VI,  1687. 

Arning  and  Klein.  Deutsch.  med.  Wochenschr.,  1907, 
XXXIII,  1482. 

Bab.     Ibid.,  1906,  XXXII,  1945. 

Bab£s  and  Mironescu.  Berlin,  klin.  Wochenschr.,  1906, 
XLIII,  1119. 

Bab£s  and  Pan6a.     Ibid.,  1905,  XLII,  1506. 

Bandi   and   Simonelli.     Riforma   Medica,    1905,    No.    29. 

Bandi  and  Simonelli.  Muenchener,  med.  Wochenschr., 
1905,  LII,  1668. 

Bandler.     Prager    med.    Wochenschr.,    1905,    XXX,    475. 

Beer.     Deutsch.   med.    Wochenschr.,    1906,   XXXII,    1192. 

Benda.     Riforma  Medica,  1906,  XXII,  1115. 

Benda.     Berlin,  klin.  Wochenschr.,  1907,  Apr.  15  and  22. 

Berger.     Muench.    med.     Wochenschr.,     1906,    LIII,    862. 

Bertarelli.     Centralbl.  f.  Bakt.,  1906,  XLI,  639. 

Bertarelli      Ibid.,  1907,  XLIII,  167,  238. 

Bertarelli  and  Volpino.*     Boll.  R.  Acad.  Med.  di  Torino, 

1905,  June  16  (ref.  Centralbl.  f.  Bakt.,  1906,  XXXVII,  230). 
Bertarelli,     Volpino     and    Bovero.     Centralbl.    f.    Bakt., 

1906,  XL,  56. 

Blaschko.     Berlin,  klin.  Wochenschr.,  1907,  XXXIV,  336. 

Bodin.     Ann.  de  Dermatol.,  1905,  984. 

Boing.     Aerztlichen  Rundschau,  1907,  No.  43. 

Bosc.     C.  R.  Soc.  de  Biol.,  1906,  LX,  338,  340. 

Bourgsdorf.*  Rousski  Vratch,  1908,  446  (abstr.  Journ. 
de  Phys.  et  Pathol.  Gin.,  1908,  769). 

Brandweiner.      Wiener  klin.  Wochenschr.,  1906,  XIX,  339. 

Bronnum  and  Ellermann.  Deutsch.  med.  Wochenschr., 
1905,  XXXI,  1757. 

Bruckner  and  Galacesco.  C.  R.  Soc.  de  Biol.,  1910,  LXVIII, 
684. 

Burnet  and  Vincent.     C.  R.  Soc.  de  Biol.,  1905,  LIX,  474. 

Buschke.*  Verein  f.  inn.  Med.,  Berlin,  Dec.  19,  1905 
(abstr.  Zentralbl.f.  inn.  Med.,  1906,  58. 

Buschke  and  Fischer.  Deutsch.  med.  Wochenschr.,  1905, 
XXXI,  791. 

Buschke  and  Fischer.  Berlin,  klin.  Wochenschr.,  1906, 
XLIII,  6,  383. 

Campbell.    Journ.  Amer.  Med.    Assoc.,    1910.    LIV,    924. 


BIBLIOGRAPHY.  I2Q 

De  Beurmann  and  Gougerot.  Bull.  Soc.  medic,  des 
Hdpitaux,  1907,  XXIV,  943. 

De  Souza  and  Pereira.  Berlin,  klin.  Wochenschr.,  1905. 
XLII,  1385. 

Dohi.     Centralbl.f.  Bakt.,  1907,  XLIV,  246. 

Doutrelepont  and  Grouven.  Deutsch.  med.  Wochenschr., 
1906,  XXXII,  908. 

Dudgeon.     Lancet,  1906,  I,  669. 

Ehrmann.     Deutsch.  med.  Wochenschr.,  1906,  No.  28. 

Erhmann.     Wiener  klin.    Wochenschr.,    1906,    XIX,    828. 

Ehrmann.     Centralbl.f.  Bakt.,  1907,  XLIV,  243. 

Ferre.     C.  R.  Soc.  de  Biol.,  1906,  LX,  97. 

Fliigel.     Deutsch.  med.   Wochenschr.,   1905,  XXXI,   1755. 

Follet.     C.  R.  Soc.  de  Biol.,  1907,  LXII,  667. 

Fouquet.     C.  R.  Acad.  des  Sciences,  1906,  CXLIII,  792. 

Fouquet.     C.  R.  Soc.  de  Biol.,  1907,  LXII,  225. 

Friedenthal.  Berlin,  klin.  Wochenschr..  1906,  XLIII, 
1217. 

Galli-Valerio  and  Lesseur.  Revue  medicale  de  la  Suisse 
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Galli-Valerio  and  Salomon.  Centralbl.  f.  Bakt..  1907, 
XLV,  37. 

Gauche  and  Merle.  C.  R.  Acad.  des  Sciences,  1909, 
CXLVIII,  862. 

Giemsa.     Deutsch.  med.  Wochenschr.,  1907,  676. 

Gierke.     Muench.  med.  Wochenschr.,  1906,  LI.II.  393. 

Gierke.     Centralbl.f.  Bakt.,  1907,  XLIV,  348. 

Gordon.*     American  Medicine,  1905,  July  23. 

Gradle.*     Trans.  Pathol.  Soc.  of  Chicago,  1907,  VII,  54. 

Greeff  and  Clausen.  Deutsch.  med.  Wochenschr.,  1906, 
XXXII,  1454. 

Grouven.     Medizinische  Klinik,  1908,  No.  8. 

Grouven  and  Fabry.  Deutsch.  med.  Wochenschr.,  1905, 
XXXI,  1469. 

Hansteen.*  Norsk.  Mag.f.  Leagevid.,  1906,  No.  7  (abstr. 
Centralbl.f.  inn.  Med.,  1906,  1193). 

Harris  and  Corbus.  Journ.  Amer.  Med.  Assoc.,  1908,  LI, 
1928. 

Harvey  and  Bousfield.  Journ.  Royal  Army  Med.  Corps, 
1905,  V,  263. 

Hedr£n.     Centralbl.f.  Bakt.,  1908,  XLVI,  232. 


130  SPIROCH^ETES. 

Herxheimer.     Deutsch.  med.  Wochenschr.,  1905.  XXX, 

i7- 

Herxheimer.     Muench.  med.   Wochenschr.,   1905,   No.   39. 
Herxheimer  and  Loser.     Ibid.,  1905,  LII,  2212. 
Herxheimer  and  Opificius.     Ibid.,  1906,  LIII,  301. 
Hoffmann.     Berlin,  klin.   Wochenschr.,   1905,  XLII,   729, 

i4Si. 

Hoffmann.     Verein.  f.  inn.    Med.,    Berlin,  1905,  Dec.  19. 
Hoffmann.     Centralbl.  f.   Bakt.,    1906,    XXXVIII,     Ref. 
(Beiheft),  108. 

Hoffmann    and     B  riming.     Deutsch.     med.     Wochenschr., 

1907,  XXXIII,  553. 

Hoffmann  and  Halle.  Muench.  med.  Wochenschr.,  1906, 
LIII,  1516. 

Hoffmann,  Lohe  and  Mulzer.      Deut.    med.    Wochenschr., 

1908,  XXXIV,  1185. 

Horand.     Lyon  midicale,  1905,  CIV,  1223. 

Huebschmann.  Berlin,  klin.  Wochenschr.,  1906,  XLIII, 
796. 

Jacquet  and  Sevin.  Bull.  Soc.  mid.  des  Hdpitauoc,  1905, 
XXII,  420. 

Jacquet  and  S£zary.     Ibid.,  XXIV,  114. 

Jancke.     Med.  Klinik,  1907,  No.  17. 

Kiolomenoglou  and  Cube.  Muench.  med.  Wochenschr., 
1905,  No.  27. 

Kowalewsky.     Deutsch.  med.  Wochenschr.,  1905,   No.  52. 

Kraus.     Prager  med.  Wochenschr.,   1906,  XXXI,  354. 

Kraus     and     Prantschoff.      Wiener     klin.      Wochenschr.. 

1905,  XVIII,  941. 

Kreibich.     Ibid.,  1906,  XIX,  199. 

Krzystalowicz  and  Siedlecki.  Bull,  de  I'Acad.  de  Sciences 
de  Cracovie,  Nov.,  1905. 

Landsteiner     and     Mucha.     Wiener     klin.     Wochenschr., 

1906,  No.  45. 

Lebailly.     C.  R.  Acad.  des  Sciences,   1908,  CXLVI,  312. 
Leixer.     Zentralbl.f.  inn.  Med.,  1905,  701. 
Leuriaux  and  v.  Geets.     Centralbl.  f.  Bakt.,   1906,  XLI 
684. 

Levaditi.     C.  R.  Soc.  de  BioL,  1905,  LVIII,  845. 

Levaditi.     Ibid.,  1905,  LIX,  326,  342. 

Levaditi.     Ann.  de  VInst.  Pasteur,  1906,  XX,  41. 


BIBLIOGRAPHY.  131 

Levaditi  and  Macintosh.     Ibid.,  1907,  XXI,  784. 

Levaditi  and  Mancunian.  C.  R.  Soc.  de  BioL,  1905, 
LIX,  527,  529. 

Lavaditi  and  Manouelian.     Ibid.,  LX,  134,  304. 

Levaditi  and  Marie.     Ibid.,  1907,  LXII,  872. 

Lavaditi  and  Salmon.     Ibid.,  1905,  LIX,  465. 

Levaditi  and  Sauvage.     Ibid.,  1905,  LIX,  344. 

Levaditi  and  Yamamouchi.     Ibid.,  1908,  LXIV,  313,  408. 

Levaditi  and  Yamamouchi.  C.  R.  Acad.  des  Sciences,  1908, 
CXLVI,  1 120. 

Lipschutz.  Deutsch.  med.  Wochenschr.,  1905,  XXXI. 
1832. 

Mclntosh.     Journ.    of  Path,   and   Bacterial.,    1909,    XIII, 

239- 

MacLennan.     Brit.  Med.  Journal,  1906,  I,  258. 

MacWeeney.     Ibid.,  1905,  I,  1262. 

Marinesco  and  Min£a.*  Revue  Neurologigue,  1906,  XIV 
388. 

Menetrier  and  Rubens-Duval.  Butt.  Soc.  mid.  des  Hdfr- 
taux,  1905,  XXII,  1059. 

Metchnikoff.     Semaine  medicale,  1905.  234. 

Methnikoff  and  Roux.  Ann.  de  I'lnst.,  Pasteur.,  1905, 
XIX,  673. 

Milman.*  Rousski  Vratch,  1906,  205  (abstr.  Journ.  de 
Phys.  et  de  Pathol.  Gin.,  1906,  556). 

Mucha  and  Scherber.  Wiener  klin.  Wochenschr.,  1906, 
XIX,  145- 

Miihlens.     Deutsch.  med.  Wochenschr.,  1907,  1207. 

Miihlens.     Centralbl.f.  Bakt.,  1907,  XLIII,  694.  586. 

Muhlens.     Ibid.,  1909,  XXXV,  1261. 

Miihlens  and   Lohe.     Centralbl.    f.    Bakt.,    1908,    XLVII, 

487. 

Miihlmann.     Ibid.,  1906,  XXXVII,  Ref.,  650. 

Mulzer.     Berlin,  klin.  Wochenschr.,  1905,  No.  36. 

Nattan-Larrier  and  Brindeau.  C.  R.  Soc.  de  BioL,  1906, 
LX,  181. 

Nicolas,  Favre  and  Andr£.     Ibid.,  1905,  LIX,  497. 

Nicolas,  Favre  and  Andr£.  Lyon  mtdicale,  1905,  LIV, 
1367. 

Niessen  (Von).  Wiener,  med.  Wochenschr.,  1908,  LVIII, 
2524,  2578. 


132  SPIROCILETES. 

Nigris.     Deutsch.  med.   Wochenschr.,    1905,  'XXXI,    1431. 

Nobdcourt,  Levaditi  and  Darr£.  C.  R.  Soc.  de  Biol.,  1905, 
LVIII,  1021. 

Noeggerath  and  Staehelin.  Muench.  med.  Wochenschr., 
1905,  1481. 

Oppenheimer  and  Sachs.  Wien.  klin.  Wochenschr.,  1905. 
XVIII,  1177. 

Pasini.     Riforma  Medica,  1907,  XXIII,  52. 

Pauli.    Johns  Hopkins  Hosp.  Bulletin,  Nov.,  1908. 

Ploeger.     Muench.  med.   Wochenschr.,  1905,  LII  1381. 

Popovitch.     Thtse  de   Paris,  1906,  No.  158. 

Preisz.*     Pester  med.-chir.  Presse,  1908,  Nov.  8  and  15. 

Proca  and  Vasilescu.  C.  R.  Soc.  de  Biol.,  1905,  LVIII, 
1044 

Qu6y.     C.  R.  Soc.  de  Biol.,  1907,  LII,  379. 

Queyrat   and   Jol  train.      Bull.    Soc.    m6d.    des.    Hdpitaux, 

i9°5.  559- 

Queyrat,  Levaditi  and  Feuillid.  Ann  de.  Dermatol.,  1905, 
982. 

Rach  and  Wiesner.     Wiener  klin.  Wochenschr,  1907,  XX, 

521- 

RadaelL     Lo  Sperimentale,  1906,  LX,  397. 

Raubitschek.     Wiener  klin.  Wochenschr.,  1905,  XVIII,  752. 

Ravaut  and  Ponselle.  Gaz.  des  Hdpitaux,  1906,  LXXIX, 
1023. 

Reischauer.     Deutsch.   med.    Wochenschr.,    1905,    XXXI, 


Renter.     Zeitschrift  f.  Hygiene,  1906,  LIV,  49. 

Ribadeau-Dumas  and  Poisot.  C.  R.  Soc.  de  Biol.,  1907, 
LXII,  247. 

Richards.     Medical  Chronicle,  1906,  X,  273. 

Richards,  and  Hunt.     Lancet,  1905,  II,  963. 

Rille.     Muench.  med.  Wochenschr.,   1905,  LII,   1377. 

Rille  and  Vockerodt.     Ibid*,  1905,  1620. 

Risso  and  Cipollina.  Riforma  medica,  1905,  XXI,  848, 
938. 

R6na.*     Wiener,  wed.  Presse,  1907,  No.  34. 

Roscher.  Berlin,  klin.  Wochenschr.,  1905,  XLII,  1382, 
1418,  1447. 

Rosenberger.  Amer.Journ.  Med.  Sciences,  1906.  CXXXI, 
143- 


BIBLIOGRAPHY.  133 

Rosenberger.*  Proc.  Pathol.  Soc.  Philadelphia,  1906,  IX, 
49;  1908,  XI,  68. 

Saling.     Centralbl.f.  Bakt.,  1906,  XLI,  737. 

Saling.     Ibid.,  1907,  XLIV. 

Saling.     Fortschritte  der  Medizin,  1907,  Hft.  19. 

Salmon.     C.  R.  Soc.  de  Biol.,  1905,  LVIII,  883. 

Sambon.     Brit.  Med.  Journ.,  1907,  II,  1321. 

Schaudinn.  Deutsch.  med.  Wochenschr.,  1905,  XXXI,  711, 
1665. 

Schaudinn  and  Hoffmann.  Berlin,  klin.  Wochenschr., 
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Schaudinn  and  Hoffmann.  Deutsch.  med.  Wochenschr., 
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Schereschewsky.     Centralbl.f.  Bakt.,  1908,  XL VII,  41. 

Schereschewsky.  Deut.  med.  Wochenschr.,  1909,  XXXV, 
835,  1260. 

Schlimpert.  Berlin,  klin.  Wochenschr.,  1906,  XXXI, 
1037. 

Schmorl.     Muench.  med.  Wochenschr.,  1907,  188,  239. 

Scholtz.     Deutsch.  med.  Wochenschr.,  1905,  XXXI,  1487, 

Schor.*  Rousski  Vratch,  1905,  1123  (abstr.  Journ.  de 
Phys.  et  Path.  Gen.,  1906,  353). 

Schuller.     Centralbl.  f.  Bakt.,  1907,  XLIII,  794. 

Schiitz.     Muench.  med.  Wochenschr.,  1906,  LIII,  543. 

Shennan.     Lancet,  1906,  I,  663,  746,  1323. 

Selenew.     Centralbl.  f.  Bakt.,  1910,  LIV,  7. 

Sezary  and Paillard.   C.  R.  Soc.  de  Biol,  1910,  LXVIII,  295. 

Siebert.     Deutsch.  med.  Wochenschr.,  1905,  XXXI,  1426. 

Siegel.     Centralbl.f.  Bakt.,  1907,  XLV,  218. 

Siegel.     Ibid.,  1906,  XL VI,  315. 

Siegel.     Ibid.,  1909,  XL VIII,  599. 

Simmonds.     Muench.  med.  Wochenschr.,  1906,  LIII,  1302. 

Sobernheim  and  Tomasczewsky.  Muenchener  med.  Woch- 
enschr., 1905,  1857. 

Spitzer.     Wien.   klin.   Wochenschr.,   1905,    XVIII,  822. 

Stanziale.     Centralbl.f.  Bakt.,  1909,  XLIX,  551. 

Thesing.     Centralbl.f.  Bakt.,  1906,  XL,  350. 

Thesing.     Muenchener  med.  Wochenschr.,  1905,  No.  28. 

Thibierge.     Gaz.  des  Hdpitauoc,  LXXIX,  123. 

Thibierge,  Ravaut.  and  Burnet.  C.  R.  Soc.  de  Biol.,  1906, 
LX,  298. 


134  SPIROCELETES. 

Thibierge,  Ravaut,  and  Le  Sourd.  Bull.  Soc.  med.  des 
Hbpitaux,  1906,  383. 

Tomasczewski.  Muench.  med.  Wochenschr.,  1906,  LIII, 
1301. 

Truffi.     Centralbl.f.  Bakt.,  1909,  XL VIII,  597. 

Uhle  and  Mackinney.*  Proc.  Pathol.  Soc.  Philadelphia, 
1906,  IX,  195. 

Uhlenhuth,  Hoffmann,  and  Weidanz.  Deutsch.  med. 
Wochenschr.,  1907,  XXXIII,  1592. 

Uhlenhuth  and  Mulzer.  Arb.  a.  d.  kaiserl,  Gesundheit- 
samte,  1909,  XXXIII,  183. 

Veillon  and  Girard.     C.  R.  Soc.  de  BioL,  1905,  LIX,  652. 

Vincent.  Bull.  Soc.  mid.  des  Hdpitaux,  1905,  XXII, 
603. 

Volpino  and  Fontana.  Centralbl.  f.  Bakt.,  1906,  XLII, 
666. 

Vuillemin.     C.  R.  Acad.  des  Sciences,   1905,  CXL,   1567. 

Wallich  and  Levaditi.     C.  R.  Soc.  de  Biol.,  1906,  LX,  191. 

Wersilowa.     Centralbl.f.  Bakt.,  1906,  XLII,  513. 

Wolff.     Ibid.,  1907,  XLIII,  803. 

Wolters.     Mediz.  klinik,  1905,  I,  963. 

Zabolotny.*  Rousski  Vratch,  1906,  212  (abstr.  Journ.  de 
Phys.  et  Pathol.  Gin.,  1906,  556). 

Zabolotny  and  Maslakowetz.  Centralbl.  f.  Bakt.,  1907. 
XLIV,  534. 

SP.    PERTENUIS. 

Asburn  and  Craig.*  Philippine  Journal  of  Science,  1907, 
II,  441- 

Castellani.*  Arch.  f.  Schiffs-  u.  Tropen-hygiene,  1907, 
XI,  19. 

Castellani.     British  Med.  Journal,  1905,  II,  1280. 

Castellani.     Deutsch.  med.  Wochenschr.,  1906,  XXXII,  132. 

Castellani.    Journ.  Trop.  Med.,  1906,  IX,  i. 

Levaditi  and  Nattan-Larrier.  Ann.  de  VInst.  Pasteur, 
1908,  XXII,  260. 

Mayer.     Deutsch.  med.  Wochenschr.,  1907,  No.  12. 

Neisser,  Baermann  and  Halstadter.  Muenchener  med. 
Wochenschr.,  No.  28. 

Schiiffner.     Muenchener    med.    Wochenschr.,     1907,    LIV, 


BIBLIOGRAPHY.  135 

Von  dem  Borne.    Journ.  of  Tropical  Med.,  1907,  X,  345. 
Wellmann.     Journ.  Trop.  Med.,  1905,  VIII,  345. 
Wellmann.*     Arch.   f.   Schiffs-  u.    Tropen-hygiene ,     1908, 
XI,  Hft.  17. 

SP.     LYMPHATICA. 

Proescher.     New  York  Med.  Journ.,  1909,  LXXXIX,  848. 

White  and  Proescher.  Journ.  Amer.  Med.  Assoc.,  1907, 
XLIX,  1115,  774. 

White  and  Proescher.  Muench.  med.  Wochenschr..  1907, 
1868. 

SP.  IN  ANEMIA. 

Moritz.  Deutsch.  Arch.  f.  klin.  Med.,  LXXXIV,  Hft. 
5-6. 

Thomas  and  Rolleston.     Brit.  Med.  Journ.,  1910,  I,  16. 

SP.    ABORIGINALIS. 

Bosanquet.     Parasitology,  1910,  II,  345. 
Cleland.     Journ.  Trop.  Med.  and  Hygiene,  1909,  May  15. 
MacLennan.     Medico-Chirurg.    Transactions,  £1907,    XC, 
700. 

Wise.     Brit.  Med.  Journ.,  1906,  I,  1274. 

SPIROCH^TA    INTERROGANS. 

Stimson.  Trans.  Soc.  Tropical  Med.,  London,  1909, 
HI,  56- 

VARIOUS    SPIROCH^TES    AND    SPIRILLA. 
ONYCHIA. 

Button,  Todd  and  Tobey.  Liverp.  Sch.  Trop.  Med.,  1906, 
Mem.  XX,  87. 

SPIROCH^TES    IN    ALIMENTARY    CANAL. 

Oppenheimer.*  Collected  Studies  Res.  Lab.  Dept.  of  Health, 
New  York,  1906,  II,  145. 

Rigaud.     C.  R.  Soc.  de  Biol.,  1909,  LXVI,  229. 
Salomon.     Centralbl.f.  Bakt.,  1896,  XIX,  433. 
Smith.     Ibid.,  1894,  XVI,  324. 
Werner.     Centralbl.f.  Bakt.,  1909,  LII,  241. 


136  SPIROCH^TES. 

SPIRILLA    IN    CHOLERA    DEJECTA. 

Abel.     Centralbl.f.  Bakt.,  1894,  XV,  213. 
Aufrecht.     Ibid.,  1894,  XVI,  405. 
Escherich.     Ibid.,  1894,  XV,  408. 
Kowalski.     Ibid.,  1894,  XVI,  321. 
Lustig  and  Giaxa.     Ibid.,  1894,  XV,  721. 
Recbtsamer.     Ibid.,  1894,  XV,  795. 

SP.  GLOSSING. 
Novy  and  Knapp.     Journ.  of  Infect.  Dis.,  1906,  III,  291. 

SPIRILLA    IN    PYELITIS    (SP.    PYOGENES). 

Mezinescu.     Centralbl.f.  Bakt.,   1904,  XXXV,   201. 

SP.    IN    DYSENTERY. 

Le  Dantec.     C.  R.  Soc.  de  Biol.,  1903,  LV,  617. 

SPIROCH^TA  DAXENSIS. 

Cantacuzfene.     Compt.      rend.      Soc.      de    Biologie,     1910, 
LXVIII,  75- 


BIBLIOGRAPHIC  INDEX. 


ABEL,  115 

Arndt,  101 

Arning  and  Klein,  50 

Ascoli,  26,  27 

Ass  my,  103 

BAB,  51,  56 

Babes  and  Pane"a,  1 1 1 

Baermann,  107 

Balfour,  44,  115 

Bandi  and  Simonelli,  29,  50,  65 

Baruchello  and  Pricolo,  26,  61, 

95 

Beer,  32,  34 
Benda,  54,  56 
Bertarelli,  56 
Bertarelli  and  Bovero,  50 
Bertarelli  and   Volpino,  30,  65 
Bertarelli  and  Volpius,  104 
Beurman  and  Gougerot,  53 
Bizzozero,  115 
Blanchard,  70,  102 
Bodin,  50 
Bonhoff,  1 06 
Borne,  112 
Borrel,  34,  91,  97 
Borrel  and  Burnet,  32 
Borrel  and  Cernovodeanu,  21 
Bosanquet,  28,  44,  114 
Bosc,  30 

Bousfield  and  Harvey,  43 
Bovero  and  Bertarelli,  50 
Branch,  30,  59,  104 
Breinl  and  Kinghorn,  62,  96 
Bruckner  and  Galacesco,  33,  53, 

in 
Briining,  65 


Burnet  and  Borrel,  32 

Burn,  66 

Butschli,  73 

Butschli  and  Schaudinn,  76 

CALKINS,  97,  98,  105 
Cantacuzene,  76,  77,  91 
Carpano  and  Martoglio,  95 
Carter,  37,  38,  40,  48,  58,  62, 

9°.  97 
Castellani,  18,  28,    29,    30,    40, 

53.  57.  59.  83,  104,  112,  113, 

114 

Cernovodeanu  and  Borrel,  21 
Certes,  77 

Cipollina  and  Risso,  50 
Cleland,  44,  58,  96,  114 
Cohn,  70,  71,  86,  100 
Collings,  76 

Compte  and  Nicolle,  34,  61,  93 
Cox,  40,  87 
Cube  and  Kiolomenoglou,  49, 

104 

DARLING,  48 

Davidsohn,  64 

De  Lisle  and  Jullien,  53 

De  Sousa  and  Pereira,  56 

Delafield,  63 

Dobell,  99 

Dodd,  62,  63,  95,  96 

Doerr,  116 

Doflein,  17,  23,  71,  73,  75 

Donne',  106 

Dudgeon,  28 

Dufouger£  and  Thiroux,  90 

Dutton,  28,  43 


137 


138 


BIBLIOGRAPHIC   INDEX. 


Button  and  Todd,  88  Grubler,  65 

Button,  Todd,  and  Tobey,  99  Gruby,  71 

Buval  and  Todd,  31,  34,  36,  89      Grunwald,  65 


EHRENBERG,  u,  70,  71,  73 
Ehrlich  and  Hata,  92 
Ehrmann,  56 
Eitner,  30,  106 
Ellermann,  59 

FANTHAM,  21,  23,  28,  34,  79,  84, 

93 

Fantham  and  Gross,  18 
Ferr6,  50 
Flournoy,  70,  87 
Flournoy,  Norris,  and  Pappen- 

heimer,  32 
Foix  and  Mallein,  64 
Foley  and  Sergent,  46 
Follet,  56    ' 

Fontana  and  Volpino,  32 
Forest,  64 
Fouquet,  30,  56 
Fraenkel,  27,  87 

GABRITCHEWSKY,  60,  88,  91 
Galacesco    and    Bruckner,    33, 

53.  I" 

Gaucher  and  Merle,  56 
Gauducheau,  44 
Gaylord,  52,  97,  105 
Geets  and  Leuriaux,  26,  32,  37, 

40,  no. 
Giemsa,  26,  63,  64,  65,  87,  98, 

102,  107,  108,  109 
Gierke,  56,  109 
Gino  de  Rossi,  64 
Girard  and  Veillon,  56 
Goadby,  101 
Goldhorn,  64 
Gonder,  20,  26,  34,  35,  36,  79, 

82,  84,  93,  94,  99,  109 
Gougerot  and  Beurman,  53 
Gross,  18,  36,  71,  86 
Gross  and  Fanthan,  18 


HAENDEL  and  Uhlenhuth,  46 

Hallopeau,  42 

Hartmann   and   Miihlens,    100, 

101 

Harvey  and  Bousfield,  43 
Hata  and  Ehrlich,  92 
Heanley,  95 
Heidenhain,  63 
Herxheimer,  28,  50 
Herxheimer  and  Hubner,  65 
Hodges  and  Ross,  89 
Hoffmann,  26,  48,  56,    101,  105 
Hoffmann  and  Prowazek,   100, 

101,  107 

Hoffmann  and  Schaudinn,  108 
Horand,  in 

Hubner  and  Herxheimer,  65 
Huebschmann,  56 
Hunt  and  Richards,  107 

JAFFE\  98 

Jancke,  56 

Jaquet  and  Sevin,  50 

Jenner,  65 

Jullien  and  Be  Lisle,  53 

KARLINSKI,  51,  69 

Kenrick,  26 

Ker,  45 

Keysselitz,i7,  21,  22,  33,  79,  80, 

81 

Kinghorn  and  Breinl,  62,  96 
Kiolomenoglou  and  Cube,   49, 

104 

Klein  and  Arning,  50 
Klodnitsky,  35,  36,  87 
Knapp  and  Novy,  86,  87,  88, 

95.  n6 

Koch,  29,  34,  36,  45,  47,  89,  101 
Kowalski,  115 
Kraus,  106 


BIBLIOGRAPHIC   INDEX. 


139 


Kraus  and  Prantschoff,  50 

Krienitz,  30,  105 

Krienitz  and  Scholtz,  49 

Krzystalowicz  and  Siedlecki, 
25.  27,  30,  34,  35,  37,  40,  41, 
42,  43,  89,  108,  109,  no,  in 

LANGE  and  Levaditi,  61,  92 

Laptschinsky,  73 

Launois  and  Loederich,  43 

Laveran,  94 

Laveran  and  Mesnil,  20,  79 

Le  Dantec,  116 

Lebailly,  32 

Lebert,  86 

Leishman,  16,  40,  43,  44 

Leixer,  56 

Leuriaux  and  von  Geets,  26,  32, 

37,  40,  no 
Levaditi,  26,  32,  39,  53,  56,  60, 

64,  65,  87,  89,  106,  107,  108, 
109,  no,  112,  113,  114 

Levaditi  and  Lange,  61,  92 
Levaditi  and  Macintosh,  109 
Levaditi   and   Mancunian,   47, 

65,  89 

Levaditi    and    Nattan-Larrier, 

S3.  57 

Levaditi  and  Stanesco,  33,  102 
Levadir.i  and  Wallich,  56 
Levaditi  and  Yamamouchi,  34 
Loederich  and  Launois,  43 
Loeffler,  66,  67 
Loewenthal,  30,  42,  101,  105 
Lucius,  65 
Liihe,  71 

MACKIE,  102 
Mallein  and  Foix,  64 
Mandelbaum,  67 
Mancunian  and    Levaditi,   47, 

65,  89 

Manteufel,  46 
Marchoux,  92 
Marchoux  and  Salimbeni,  91 


Marshall,  55 
Martin,  35,  60,  95 
Martoglio  and  Carpano,  95 
Maskalowetz   and  Zabolotony, 

56 

Mayer,  28,  39,  42,  43,  89 
Mayer  and  Newstead,  34 
Mclntosh,  57,  112 
Mclntosh  and  Levaditi,  109 
McKinnon,  34,  88 
McLennan,  58 
McNeal,  64 
McNeal  and  Novy,  16 
Meirowsky,  67 
Meister,  65 

Merle  and  Gaucher,  56 
Mesnil  and  Laveran,  20,  79 
Metchnikoff,  46,  50,  70 
Mezinescu,  116 
Migula,  70 
Miller,  29,  101 
Milne  and  Ross,  88 
Minchin,  68 
Moebius,  77 
Moffat,  89 
Moritz,  30,  59,  113 
Mtihlens,  50,  101,  in 
Miihlens   and   Hartmann,    100, 

IOI 

Miihlens  and  Schereschewsky,32 
Mulzer,  104 

NAPLES,  75 

Nattan-Larrier    and    Levaditi, 

S3,  57 

Neufeld  and  Prowazek,  60 
Neumann,  98,  99 
Newstead  and  Mayer,  34 
Nicolle,  98 

Nicolle  and  Compte,  34,  61,  93 
Niessen,  42 
Norris,  70,  87 
Norris,  Pappenheimer,  and 

Flournoy,  32 
Novy,  48,  114 


140 


BIBLIOGRAPHIC   INDEX. 


Novy  and  Knapp,  86,  87,  88,  95, 

116 

Novy  and  MacNeal,  16 
Nuttall,  34,  88 

OBERMEIER,  45 
Oppenheimer,  115 
Oppenheimer  and  Sachs,  64 

PANE  A  and  Babes,  in 
Pappenheimer,  70,  87 
Pappenheimer,  Norris,  and 

Flournoy,  32 
Pereira  and  De  Sousa,  56 
Perrin,  17,  18,  19,  20,  21,  22,  23, 

33,  36,  37,  38,  40,  77,  78,  79, 

80,  84 
Petrie,  44 
Plaut,  25,  76,  1 02 
Polland,  103 

Ponselle  and  Ravaut,  66 
Popovitch,  87 
Porter,  34 

Prantschoff  and  Kraus,  50 
Pricolo  and  Baruchello,  26,  61, 

95 

Proca  and  Vasilescu,  64 
Proescher,  113 

Proescher  and  White,  57,  113 
Prowazek,  26,  27,  29,  34,  35,  38, 

41,  42,  84,  91,  92,  94,  98,  101, 

103 
Prowazek    and   Hoffmann,   48, 

100,  101,  107 
Prowazek  and  Neufeld,  60 

QUE"RY,  42 

RABINOWITSCH,  46 
Ravaut  and  Ponselle,  66 
Rechtsamer,  115 
Reichert,  27 
Reitmann,  64 
Reuter,  29,  56 
Richards  and  Hunt,  107 


Rigaud,  115 

Rille,  1 06 

Rille  and  Vocquerodt,  54 

Risso  and  Cipollina,  50 

Romanowsky,  65 

R6na,  104 

Ross  and  Hodges,  89 

Ross  and  Milne,  88 

Rossi,  98 

SACHAROFF,  90,  91 

Sachs  and  Oppenheimer,  64 

Sakurane,  105 

Salimbeni  and  Marchoux,  91 

Saling,  68,  69 

Salomon,  115 

Sambon,  108 

Schaudinn,  n,  16,  26,  27,  28, 
48,  53,  69,  70,  71,  74,  75,  100, 
105,  106,  108,  109,  114 

Schaudinn  and  Biitschli,  76 

Schaudinn  and  Hoffmann,  108 

Schaudinn  and  Vuillemin,  25 

Schellack,  18,  20,  21,  27,  28,  30, 
33.  34,  35,  38,  46,  47,  78,  80, 
81,  82,  83,  84,  85,  87,  89 

Scherber,  107 

Schereschewsky,  30,  33,  43,  63, 

102,    IO9,    III 

Schereschewsky    and    Muhlens 

32 

Schmeidlechner,  14,  43 

Schmorl,  56,  64 

Scholtz  and  Krienitz,  49 

Schrdtter,  70 

Schiiffner,  53,  57,  112 

Schwellengrebel,  12,  13,  33 

Sergent,  16 

Sergent  and  Foley,  46 

Sevin  and  Jaquet,  50 

Shennan,  50 

Siebert,  50 

Siedlecki  and  Krzystalowicz, 
25,  27,  30,  34,  35,  37,  40,  41, 
42,  43,  89,  108,  109,  no,  in 


BIBLIOGRAPHIC   INDEX. 


141 


Siegel,  no 

Simmonds,  56 

Simonelli  and  Bandi,  29,  50,  65 

Smith  (T.)  116 

Sobernheim  and  Tomasoli,  30 

Stanesco  and  Levaditi,  33,  102 

Stephens,  26,  27 

Stimson,  114 

Stordy,  61 

THEILER,  44,  61,  94,  95 

Thiroux  and  Dufougere,  90 

Tictin,  46 

Tobey  and  Todd,  28,  43 

Tobey,  Dutton,  and  Todd,  99 

Todd  and  Dutton,  88 

Todd  and  Duval,  31,  34,  36,  89 

Todd  and  Tobey,  28,  43 

Todd,  Dutton,  and  Tobey,  99 

Tomasoli    and  Sobernheim,  30 

Tunnicliffe,  43 

UFFENHEIMER,  59 
Uhlenhuth  and  Haendel,  46 

VASILESCU  and  Proca,  64 
Veillon  and  Girard,  56 
Veszpremi,  39,  59,  102 
Vincent,  102 

Vocquerodt  and  Rille,  54 
Volpino  and  Bertarelli,  30,  65 


Volpino  and  Fontana,  32 
Volpius  and  Bertarelli,  104 
von  dem  Borne,  57,  65 
von  Geets  and  Leuriaux,  26,  32, 

37,  40,  no 
von  Niessen,  42 
Vuillemin,  69,  71,  108 
Vuillemin  and  Schaudinn,  2  5 

WALLICH  and  Levaditi,  56 

Wassermann,  56 

Waters,  59,  104 

Wellmann,  43,  53,  57,  113 

Wenyon,  35,  44,  62,  96,  97 

Werner,  116 

White  and  Proescher,  57,  113 

Whitmore,  103 

Wibel,  70 

Williams,  32 

Williams  (L.  A.  and  R.  S.)  89 

Wise,  58,  114 

Wolff,  86 

YAMAMOTO,  66 

Yamamouchi  and  Levaditi,  34 

ZABOLOTNY  and  Maskalowetz, 

56 

Zettnow,  29,  92 
Ziehl,  64 
Zuelzer,  74 


INDEX. 


AFRICAN  tick  fever,  46 
Alimentary  canal  of   dog,    spi- 
rilla in,  115 
spirochaetes  in,  115 
spirochaetes  in,  116 
American  relapsing  fever,  47 
Angina,  Vincent's,  58 
Animals,  diseases  of,  60 
Antisyphilitic  treatment,  effect 
of,   on  spirochaeta  pallida,  54 
Arabian    spirochsete,    conjuga- 
tion in,  38 
sporulation  in,  40 
Association  of  spirochaetes  with 

other  organisms,  42 
Atoxyl  in  fowl  spirochaetosis,  61 
Autogamy,  79 
Axenstrang,  92 

BACILLUS    fusiformis,    associa- 
tion of  spirochaetes  with, 

43 

spirochaeta    duttoni    asso- 
ciated with,  43 
pallida   associated  with, 

43 
pertenuis     associated 

with,  43 
refringens  associated 

with,  43 
Balanitis,  60 
Basalkorn,  80 
Bat,  disease  due  to  spirochaete 

in,  6 1 
Bertarelli  and  Volpino's  stain, 

65 
Bibliography,  117-136 


Biological  position,  1 1 
Blanchard  's    classification,     70 
Blepharoplasts,  20 
Blood-spirochaetes,  71 
Bodies,  coccoid,  associated  with 
spirochaeta  anodontae,  41 
ovoid,  40 

rod-shaped,    associated   with 
spirochaeta  duttoni,  42 
gallinarum,  42 
lutrae,  42 
pallida,  41 
with  spirochaetes,  41 
Bombay,  relapsing  fever  in,  48 
Bronchitis,  59 

haemorrhagic,  59 
Burn's    india   ink    method    of 
identifying  spirochastes,  66 

CALCULOUS  pyelitis,  pus  from, 

spirilla  in,  1 1 6 

Canal,  alimentary,  of  dog,  spi- 
rilla in,  115 
spirochaetes  in,  115 
spirochaetes  in,  116 
Cancer  in  mice,  spirochastes  in, 

52,  97 
Cattle,  fever  of,  spirochaetes  in, 

61 
Chancre,  spirochaeta  pallida  in, 

5i 
Chemical   reagents,    action   of, 

on  small  spirochastes,  15 
Cholera,  feces  of,  spirilla  in,  115 
Chromatin  in  spirochaeta  ano- 
dontae, 21 

in  spirochaeta  balbianii,  21 


143 


144 


INDEX. 


Chromatinbrocken,  94 
Classification,  69 
Coccoid  bodies  associated  with 
spirochaeta  anodontae,  41 
forms   from  spirochaeta  dut- 

toni,  40 

Congenital  syphilis,  56 
Conjugation,  14,  37 

in  spirochaeta  anodontae,  3  7 
Arabia,  38 
balbianii,  37 
pallida,  37 

Corpuscles,  pebrine,  94 
Cristispira,  71 

interrogationis,  86 
pectinis,  86 
Cultivation,  31 
failure  of,  13 
of   spirochaeta   balanitidis, 

33 

duttoni,  31 
gallinarum,  32 
gracilis,  33 
obermeieri,  32 
pallida,  32,  33,  53,  in 
refringens,  32 

DARK  field  examination,  66 
Davidsohn's  stain,  64 
Degeneration,   moniliform,    29, 

40 
Description  of  species,  72 

of  spirilla,  86 
Development,  33 

into  filaments,  36 
Division  by  incurvation,  36 

in    spirochaeta     anodontae, 

33 

balbianii,  33 
duttoni,  34 
gallinarum,  34 
hartmanni,  35 
obermeieri,  34 
pallida,  34 
vesperuginis,  34 


Division,  longitudinal,  in  large 

spirochaetes,  14 
in  small  spirochaetes,  14 
mode  of,  33 

Dog,  alimentary  canal  of,  spi- 
rilla in,  115 
spirochsetes  in,  115 
Dysentery,  feces  of,  spirilla  in, 
116 

ENCYSTMENT  in  spirochaeta  ano- 
dontae, 39 
balbianii,  38 
duttoni,  39 
pallida,  39 
Ends  of  large  spirochaetes,  shape 

of,  1 8 

Enlargement  of  spleen  in  mice 
infected  with  spirochaeta 
laverani,  62 
Entoplasm,  19 

structure  of,  21 

Euglenoid  movement  in  spiro- 
chaeta vincenti,  25 
Examination    of    living    spiro- 
chaetes, 66 
dark  field  method,  66 

FECES  of  cholera,  spirilla  in,  115 

of  dysentery,  spirilla  in,  116 
Fetus,  spirochaeta  pallida  in,  56 
Fever,  African  tick,  46 
of  cattle,  spirochaetes  in,  61 
of  horses,  spirochaetes  in,  6 1 
of  sheep,  spirochaetes  in,  61 
relapsing,  45 
American,  47 
in  Bombay,  48 
in  Panama,  48 
Filaments,      development      of 

spirochaetes  into,  36 
in  spirochaeta  duttoni,  36 

obermeieri,  36 

Fish,  shell-,  spirochaetes  in,  82, 
83 


INDEX. 


145 


Flagella  in  spirochaeta  duttoni, 

27 

obermeieri,  27 
of  large  spirochaetes,  14 
Fly,  tse-tse,  stomach  of,  spirilla 

in,  1  1  6 

Forest's  stain,  64 
Form,  change  of,  in  spirochseta 

pallida,  30 
variability  of,  29 

of  spirochaeta  pallida,  49 
Fowls,  spirillosis  of,  60 
Framboesia  tropica,  57 
Fusiform  bacillus.     See  Bacillus 
fusiformis. 

GANGRENE,  spirochaetes  in,  103 
Gangrenous    stomatitis,    spiro- 

chsetes  in,  44 
Gedrungen,  114 
Giemsa's  stain,  63 
Gino  de  Rossi's  stain,  64 
Goldhorn's  stain,  64 
Granules  in  spirochaetes,  27 
Granuloma,   ulcerative,   of  pu- 

denda, 58 
spirochaetes  in,  44 
spirochaetes     resembling 

spirochaeta  pallida  in, 

52 

Grunwald's  stain,  65 
Gumma,  spirochaeta  pallida  in, 


HABITAT,  31 

Haemorrhagic  bronchitis,  59 

Horses,  fever  of,  spirochaetes  in, 

61 
Hospital  gangrene,  spirochaetes 

in,  103 

IMMOBILISINES,  61 
Incurvation,  division  by,  36 
India    ink    method    of    identi- 
fying spirochaetes,  66 


Intestines  of  pig,  spirilla  in,  1 16 
Intra-vitam  staining,  67 
Involution  forms,  24 

KARYOSOME  in  spirochaeta  bal- 

bianii,  21 
Kriechend,  74 

LARGE    spirochaetes,     73.     See 

also  Spiroch&tes,  large. 
Leg,  tropical  ulcer  of,  103 
Length    of    large    spirochaetes, 

18 
Levaditi       and      Mancunian's 

stain,  65 

Levaditi's  stain,  65 
Lice  in  conveying  relapsing 

fever,  46 
Living     spirochaeta     dentium, 

staining,  67 
pallida,  staining,  67 
spirochaetes,  examination  of, 

66 

dark  field  method,  66 
staining,  67 
Longitudinal    division  in  large 

spirochaetes,  14 
in  small  spirochaetes,  14 
Loop  in  spirochaeta  pallida,  29 

pertenuis,  29 
in  spirochaetes,  29 
Luhe's  division  of  spirochaetes, 

7i 
Lymph,     vaccine,    spirochaetes 

in,  1 06 

Lymphadenoma,  57 
Lymphosarcoma,  57 

MALIGNANT  syphilis,  50,  55 
Mandelbaum's  stain,  67 
Mancunian    and     Levaditi's 

stain,  65 

Masses,  tangled,  29 
McNeal's  stain,  64 
Meirowsky's  stain,  67 


146 


INDEX. 


Membrane,  undulating,  in  spiro- 

chaeta  buccalis,  26 
gallinarum,  26 
refringens,  26 
vesperuginis,  26 
of  large  spirochaetes,  14, 

20 
of  small  spirochastes,  14, 

26 
Mice,  cancer  in,  spirochaetes  in, 

52,  97 

infected  with  spirochaeta 
laverani,  enlargement  of 
spleen  in,  62 

Molluscs,  spirochaetes  of,  82 
Moniliform  degeneration,  29,  40 
Morphology,  16 

Movements,  euglenoid,  in  spiro- 
chaeta vincenti,  25 
in  spirochaeta  pallida,  25 
in  treponema  pallidum,  25 
of  large  spirochastes,  19 
of  small  spirochaetes,  25 
Mouse-cancer,    spirochaetes    in, 

52.  97 

Multiplication,  33 
Mussels,  spirochaetes  from,  85 

NODULES  in  spirochaeta  aborig- 

inalis,  28 
duttoni,  28 
pallida,  28 
pertenuis,  28 
in  spirochastes,  27 
Noma,  spirochaetes  in,  104 
Nucleus     in     spirochaeta     bal- 
bianii,  21 

ONYCHIA,  spirochastes  in,  43 
Oppenheimer  and  Sachs'  stain, 

64 

Ornithodorus  moubata,  46 
Ovoid  bodies,  40 

in  spirochaeta  pallida,  40 
pertenuis,  40 


PANAMA,  relapsing  fever  in,  48 

Pathogenicity,  45 
summary  of,  62 

P6brine  corpuscles,  94 

Pediculi  capitis  et  corporis  in 
conveying  relapsing  fever,  46 

Periplast,  19 

Periplastfortstaze,  80 

Phagedaenic  ulcers,  103 

Phagocytosis,  effect  of,  on  spiro- 
chaeta pallida,  56 

Pian,  57 

Pig,  intestines  of,  spirilla  in,  1 1 6 
skin   of,  lesions,  spirochaetes 

in,  62 
spirochaetes  in,  95 

Ponselle  and  Ravaut's  stain,  66 

Pseudo-spirochaetes,  67 

Pudenda,  ulcerative  granuloma 

of,  58 

spirochaetes  in,  44 
spirochaetes     resembling 
spirochasta   pallida  in, 
52 

Pulmonary  tuberculosis,  spiro- 
chaetes in,  59 

Pus  from  calculous  pyelitis,  spi- 
rilla in,  116 

Pyelitis,    calculous,    pus    from, 
spirilla  in,  1 1 6 

RANDFIBRILLE,  20 

Ravaut    arid    Ponselle's    stain, 

66 
Reaction,    Wassermann,   in 

syphilis,  56 
Recurrent  fever,  spirochaetes  of, 

12 

Reitmann's  stain,  64 
Relapsing  fever,  45 
'American,  47 
in  Bombay,  48 
in  Panama,  48 

Relation    between    large    and 
small  spirochaetes,  u 


INDEX. 


147 


Reproduction    in    large    spiro- 

chaetes,  14 

in  small  spirochaetes,  14 
Rod-shaped   bodies    associated 
with  spirochaeta  dut- 
toni,  42 
gallinarum,  42 
lutrae,  42 
pallida,  41 
with  spirochaetes,  41 

SACK  and  Oppenheimer's  stain, 

64 
Salvarsan  (606)  in  fowl  spiro- 

chaetosis,  61 
Schellack's  list  of  spirochaetes 

found  in  shell-fish,  83 
Schereshewsky's  stain,  63 
Schmorl's  stain,  64 
Serum,  syphilitic,  action  of,  on 

spirochaeta  pallida,  53 
Sexual     cycle     of     spirochaeta 

pallida,  no 

Shape  of  large  spirochaetes,  18 
of  small  spirochaetes,  25 
of  spirochasta  anodontae,  18 
balbianii,  18 
plicatilis,  18 
Sheath  in  spirochaeta  buccalis, 

26 

duttoni,  26 
pallida,  26 
Sheep,  fever  of,  spirochaetes  in, 

61 
Shell-fish,    spirochaetes   in,    82, 

83 

Silver  stain,  65 

Skin  of  pig,   lesions  of,  spiro- 
chaetes in,  62 
Small    spirochaetes,     86.      See 

also  Spirochates,  small. 
Small-pox,  spirochaetes  in,  105 
Species,  description  of,  72 
Spirilla,  description  of,  86 
doubtful,  115 


Spirilla  in  alimentary  canal  of 

dog,  115 
in  feces  of  cholera,  115 

of  dysentery,  116 
in  intestines  of  pig,  1 1 6 
in   pus   from  calculous  pye- 

litis,  116 

in  stomach  of  tse-tse  fly,  116 
Spirillicidins,  46 
Spirillosis  of  fowls,  60 
Spirillum,  71 

giganteum,  12,  13 
glossinae,    116 
minor,  97 
obermeieri,  12 
sputigenum,  101 
volutans,  12 
Spirobacillus,  70 
Spirobacteria,  70 
Spirochaeta,  n,  71 
aboriginalis,  114 

nodules  in,  28 
acuminata,  113 
anodontae,  12,  14,  79 

chromatin  in,  21 

coccoid      bodies       associ- 
ated with,  41 

conjugation  in,  37 

description,  79 

division  in,  33 

encystment  in,  39 

shape  of,  18 

sporulation  in,  40 

staining  material  in,  24 

structure  of,  17,  21 
anserina,  90 

description,  90 
Arabia,  conjugation  in,  38 

sporulation  in,  40 
balanitidis,  107 

cultivation,  33 

description,  107 
balbianii,  12,  13,  14,  77 

chromatin,  in  21 

conjugation  in,  37 


148 


INDEX. 


Spirochaeta    balbianii,    descrip- 
tion, 77 

division  in,  33 

encystment  in,  38 

female  forms,  78 

karyosome  in,  2 1 

male  forms,  79 

nucleus  in,  21 

shape  of,  18 

sporulation  in,  40 

structure,  16,  21 
bo  vis,  95 

bronchi alis,  59,  104 
buccalis,  100 

description,  100 

sheath  in,  26 

undulating   membrane 

in,  26 
bufonis,  99 
carteri,  88 
culicis,  98 
daxensis,  76 

description,  76 

in  water,  31 
denticola,  101 
dentium,  101 

description,  101 

living,  staining  of,  67 
duttoni,  88 

associated  with  bacillus 
fusiformis,  43 

coccoid  forms  from,  40 

cultivation,  31 

description,  88 

division  in,  34 

encystment  in,  39 

filaments  in,  36 

flagella  in,  27 

nodules  in,  28 

rod-shaped     bodies     asso- 
ciated with,  42 

sheath  in,  26 
equi,  95 

eurogyrata,  116 
gadi,  98 


Spirochaeta  gallinarum,  60,  91 

cultivation,  32 

description,  91 

division  in,  34 

rod-shaped     bodies     asso- 
ciated with,    42 

undulating  membrane  in,  2  6 
gondii,  98 
gracilis,  102 

cultivation,  33 

description,  102 
hartmanni,  99 

description,  99 

division  in,  35 
interrogans,  114 
interrogationis,  86 
jonesii,  99 
lagopodis,  93 
laverani,  96 

enlargement   of   spleen  in 

mice  infected  with,  62 
lovati,  93 
lutrae,  94 

rod-shaped     bodies     asso- 
ciated with,  42 
lymphatica,  57,  113 
mactrae,  84 
media,  101 
microgyrata,  105 
minei,  98 
minor,  97 
muris,  62,  96 
novyi,  47,  88 
obermeieri,  12,  45,  86 

cultivation,  32 

description,  86 

division  in,  34 

filaments  in,  36 

flagella  in,  2  7 

staining,  87 
obtusa,  113 
ovis,  95 
pallida,  48,  108 

action  of  syphilitic  serum 
on,  53 


INDEX. 


I49 


Spirochaeta    pallida    associated 
with  bacillus  fusiformis, 

43 

change  of  form  in,  30 

conjugation  in,  37 

cultivation,  32,  33,  53,  in 

description,  108 

developmental  form  of  a 
bacillus,  42 

division  in,  34 

effect  of  anisyphilitic  treat- 
ment on,  54 
of  phagocytosis  on,  56 
of  syphilitic  virus  on,  54 

encystment  in,  39 

in  chancre,  51 

in  congenital  syphilis,  56 

in  fetus,  56 

in  gumma,  51 

in  lesions  artificially  pro- 
duced by  inoculation  of 
lower  animals  with 
spyhilis,  52 

in  non-syphilitic  condi- 
tions, 49 

living,  staining  of,  67 

loop  in,  29 

movement  in,  25 

nodules  in,  28 

objections  to,  as  cause  of 
syphilis,  49 

ovoid  bodies  in,  40 

relationship  of,  to  actual 
lesions  of  syphilis,  50 

rod-shaped  bodies  asso- 
ciated with,  41 

sexual  cycle,  no 

sheath  in,  2  6 

spirochaetes  resembling, 
in  ulcerative  granu- 
loma  of  pudenda,  52 
in  yaws,  52 

staining,  63,  64 

variability  of  form,  49 
pallidula,  112 


Spirochaeta  pectinis,  86 
pelamydis,  99 
pertenuis,  53,  57,  112 

associated     with     bacillus 
fusiformis,  43 

description,  112 

loop  in,  29 

nodule  in,  28 

ovoid  bodies  in,  40 

pinnae,  79,  84 

description,  84 
pitheci,  90 
plicatilis,  n,  73 

description,  73 

in  water,  31 

shape  of,  18 

structure,  17,  23 
polyspira,  31,  86 
pseudopallida,  104 
recurrentis,  12,  86 
refringens,  106 

associated     with     bacillus 
fusiformis,  43 

cultivation,  32 

description,  106 

undulating   membrane  in, 

26 

schaudinni,  103 
spiculifera,  19,  8 1 
stenogyrata,  116 
suis,  95 
tapetos,  84 
theileri,  94 

description,  94 
vesperuginis,  61,  93 

description,  93 

division  in,  34 

undulating   membrane  in, 

26 
vincenti,  102 

description,  102 

euglenoid     movement    in, 

25 
Spirochaetes,   associated  forms, 

3° 


INDEX. 


Spirochaetes,  association  of,  with 
bacillus  fusiformis,  43 

with  other  organisms,  42 
biological  position,  1 1 
blood-,  71 
change  in  form,  29 
classification  of,  69 
conjugation  in,  14,  37 
cultivation  of,  31 

failure,  13 
development  of,  33 

into  filaments,  36 
division  in,  33 

by  incurvation,  36 
doubtful,  115 
entoplasm  of,  19 

structure,  21 

form  of,  variability  in,  29 
general  characters,  n 
granules  in,  27 
habitat  of,  31 
in  alimentary  canal,  1 1 6 

of  dog,  115 

in  cancer  in  mice,  52,  97 
in  congenital  syphilis,  56 
in  fever  of  cattle,  61 

of  horses,  61 

of  sheep,  61 
in  gangrenous  processes,  103 

stomatitis,  44 
in  lesions  of  skin  in  pig,  62 
in  mouse-cancer,  52,  97 
in  noma,  104 
in  onychia,  43 
in  phagedaenic  ulcer,  103 

in  Pig,  95 

in  shell-fish,  82,  83 

in  small-pox,  105 

in  stomach,  change  in  form, 

30 

in  tropical  ulcer  of  leg,  1 03 
in  ulcerative  granuloma,  44 
in  vaccine  lymph,  106 
india   ink    method   of    iden- 
tifying, 66 


Spirochaetes,  large,  73 

and    small,     relation     be- 
tween, ii 

ends  of,  shape,  18 

flagella  of,  14 

length  of,  1 8 

longitudinal  division,  14 

movements  of,  19 

reproduction  in,  14 

shape  of,  18 

staining    material    in,    ar- 
rangement of,  13 

structure  of,  16,  19 

undulating  membrane   of, 

13,  14,  20 

Irving,  examination  of,  66 
dark  field  method,   66 

staining  of,  67 
loop  in,  29 

masses      of,      tangled       to- 
gether, 29 
moniliform  degeneration,  29, 

40 

morphology  of,  16 
multiplication  of,  33 
nodules  in,  27 
objects  resembling,  69 
of  African  tick  fever,  46 
of  balanitis,  60 
of  bronchitis,  59 
of  haemorrhagic  bronchitis,  59 
of  lymphadenoma,  57 
of  molluscs,  82 
of  mussels,  85 

of  pulmonary  tuberculosis,  59 
of  relapsing  fever,  45 
American  form,  47 
in  Bombay,  48 
in  Panama,  48 
of  spirillosis  of  fowls,  60 
of  syphilis,  48 
of  ulcer  of  stomach,  59 
of  ulcerative  granuloma,  58 
of  Vincent's  angina,  58 
of  yaws,  57 


INDEX. 


Spirochaetes,  ovoid  bodies  in,  40 
pathogenicity  of,  45 

summary,  62 
periplast  of,  19 
pseudo-,  67 

resembling    spirochaeta    pal- 
lida     in    ulcerative 
granuloma    of     pu- 
denda, 52 
in  yaws,  52 

rod-shaped     bodies     associ- 
ated with,  41 
small,    action    of      chemical 

reagents  on,  15 
and     large,     relation     be- 
tween, ii 
description  of,  86 
movements  of,  25 
nature  of,  14 
reproduction  in,  14 
shape  of,  25 
structure  of,  25,  26 
undulating    membrane   of, 

14,  26 

staining  of,  methods,  63 
trypanosomes    associated 

with,  44 

variability  in  form,  29 
Spironema,  69 
pallida,  108 

Spiroschaudinna  pallida,  108 
Spirosoma,  70 

Spleen,  enlargement  of,  in  mice 
infected      with      spirochaeta 
laverani,  62 
Sporulation,  40 

in  spirochaeta  anodontae,  40 
Arabia,  40 
balbianii,  40 
Stain,  Bertarelli  and  Volpino  's, 

65 

Burri's,  66 
Davidsohn's,  64 
Forest's,  64 
Giemsa's,  63 


Stain,  Gino  de  Rossi's,  64 
Goldhorn's,  64 
Grunwald's,  65 
Levaditi  and  Mancunian's,  65 
Levaditi's,  65 
Mandelbaum 's,  67 
McNeal's,  64 
Meirowsky's,  67 
Ravaut  and  Ponselle's,  66 
Reitmann's,  64 
Schereshewsky's,  63 
Schmorl's,  64 
silver,  65 
Yamamoto's,  66 
Staining,  intra-vitam,  67 

living  spirochaeta  dentium,  67 

pallida,  67 
Spirochaetes,  67 
material  in  large  Spirochaetes, 

arrangement  of,  13 
in  spirochaeta  anodontae,  24 
methods  of,  63 
spirochaeta  obermeieri,  87 

pallida,  63,  64 
Stomach  of  tse-tse  fly,  spirilla 

in,  116 
Spirochaetes     in,     change    in 

form,  30 
ulcer  of,  59 
Stomatitis,   gangrenous,   spiro- 

chaetes  in,  44 
Structure  of  large  Spirochaetes, 

1 6,  19 

of  small  Spirochaetes,  25,  26 
of  spirochaeta  anodontae,   17, 

21 

balbianii,  16,  21 

plicatilis,  17,  23 
Syphilis,  48 

clinical  phenomena,  54 
congenital,  56 
malignant,  50,  55 
relationship     of     spirochaeta 

pallida    to   actual    lesions 

of,  50 


152 


INDEX. 


Syphilis,  treatment  of,  effect  on 

spirochasta  pallida,  54 
Wassermann  reaction  in,   56 
Syphilitic  serum,  action  of,  on 

spirochaeta  pallida,  53 
virus,    effect    of,     on    spiro- 
chaeta pallida,  54 

TANGLED  masses  of  spirochaetes, 

29 

Tastend,  74 
Tick  fever,  African,  46 
Treponema,  70,  71 

pallidum,  108 

movements  in,  25 
Tropical  ulcer  of  leg,  103 
Trypanosoma,  71 

balbianii,  77 

dimorphon,  44 

luis,  37,  no 
Trypanosomes  associated  with 

spirochaetes,  44 
Trypanosomidae,  71 
Tse-tse  fly,  spirilla  in  stomach 

of,  116 

Tuberculosis,  pulmonary,  spiro- 
chaetes in,  59 

ULCER,  gangrenous,  spirochaetes 

in,  104 

of  stomach,  59 
phagedaenic,  103 
tropical,  of  leg,  103 
Ulcerative    granuloma  of  pu- 
denda, 58 


Ulcerative  granuloma  of  pu- 
denda, spirochagtes  in, 
44 

spirochaates  resembling 
spirochaeta  pallida  in, 
52 

Undulating  membrane  in  spiro- 
chaeta buccalis,  26 
gallinarum,  26 
refringens,  26 
vesperuginis,  26 
of  large  spirochaetes,  14,  20 
of  small  spirochaetes,  14,  26 

VACCINE    lymph,     spirochaetes 
in,  106 

Variability    of    form    of    spiro- 
chaeta pallida,  49 

Variola,  spirochaetes  in,  105 

Vesperugo  kuhlii,  61 

Vibrio,  70 
lineola,  106 

Vincent's  angina,  58 

Volpino  and  Bertarelli's  stain, 
65 

WASSERMANN  reaction  in  syphi- 
lis, 56 

Water,  spirochaeta  daxensis  in, 31 
plicatilis  in,  31 

YAMAMOTO'S  stain,  66 
Yaws,  57 

spirochaetes  resembling  spi- 
rochaeta pallida  in,  52 


Date  Due 


.S.A.  CAT.   NO.   24    161 


A  000  499  571  8 


QW155 


1911 
Bosancuet,  William  Cecil. 

Spirochaetes  ;  a  review  of  recent 
work... 


MEDICAL  SCIENCES  LIBRARY 

UNIVERSITY  OF  CALIFORNIA,  IRVINE 

IRVINE,  CALIFORNIA  92664 


